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Copyright © 2007 New Age International (P) Ltd., Publishers
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Preface ............................................................................................................... (v)
Chapter 1 Cell Biology ................................................................................................. 1–60
• General Introduction................................................................................................ 1
• Cell membrane ........................................................................................................ 4
• Structural organization of a cell ................................................................................ 6
• Cell cycle ............................................................................................................. 12
• Cell reproduction and Cell division .......................................................................... 15
• Cell differentiation and Cell-Cell interaction .............................................................. 23
• Cancer and Malignant growth ................................................................................. 24
– Characteristics of cancer cell ..................................................................... 25
– Types of cancer ........................................................................................ 25
– Protooncogenes and tumor suppressor genes ............................................... 26
• Immune response.................................................................................................. 27
– Innate immunity........................................................................................ 27
– Adaptive immunity .................................................................................... 28
• Humoral and cell mediated immune response............................................................ 30
• Vaccines ............................................................................................................... 31
• Dosage compensation ............................................................................................ 34
• Sex determination. ................................................................................................. 35
• Practice Test Paper-I ............................................................................................. 37
• Practice Test Paper-II ............................................................................................ 48
Chapter 2 Biochemistry ........................................................................................... 61–139
• Basic Chemistry .................................................................................................... 61
– Properties of water ................................................................................... 61
– Defination of pH and pOH.......................................................................... 61
– Buffers .................................................................................................... 62
Contents Contents
viii CSIR-NET Life Sciences
– Atoms ...................................................................................................... 62
– Molecules and chemical bonding ................................................................ 63
• Bioenergetics ........................................................................................................ 65
• Enzymes .............................................................................................................. 66
– Enzyme kinetics ........................................................................................ 70
– Control of enzyme activity ......................................................................... 71
– Mechanism of action ................................................................................. 73
• Lipids .................................................................................................................. 74
• Carbohydrates ...................................................................................................... 79
• Sugars ................................................................................................................. 82
• Glycolysis ............................................................................................................ 83
• Amino acids ......................................................................................................... 90
• Protein Structure and folding.................................................................................. 93
• DNA structure ...................................................................................................... 98
• DNA replication .................................................................................................. 100
• DNA Repair ........................................................................................................ 101
• RNA structure, Transcription and RNA Processing ................................................ 103
• Proteins synthesis ............................................................................................... 106
• Metabolic interrelationship ..................................................................................... 111
• Practice Test Paper-I ............................................................................................ 114
• Practice Test Paper-II .......................................................................................... 127
Chapter 3 Physiology ............................................................................................. 140–186
• Response to stress .............................................................................................. 140
– Water stress ........................................................................................... 140
– Oxygen stress ........................................................................................ 140
– Salt stress .............................................................................................. 141
– Heat stress ............................................................................................. 141
– Cold stress ............................................................................................. 141
• Transport across membrane ................................................................................. 142
• Plant Hormones .................................................................................................. 144
– Auxin ..................................................................................................... 144
– Gibberellin .............................................................................................. 145
– Cytokinin ............................................................................................... 147
– Ethylene ................................................................................................. 148
– Abscisic acid .......................................................................................... 148
• Animal Hormones ................................................................................................ 149
• Summary of endocrine glands and their harmones .................................................. 151
• Reproduction ...................................................................................................... 157
Contents ix
• Gametoenesis ..................................................................................................... 159
• Fertilization in humans ......................................................................................... 161
• Practice Test Paper-I ........................................................................................... 162
• Practice Test Paper-II .......................................................................................... 176
Chapter 4 Genetics ................................................................................................ 187–248
• The Foundation of Genetics ................................................................................. 187
• Mendels experiments and the laws of inheritence.................................................... 188
• Some Exceptions to Mendels laws ........................................................................ 193
• Alleles and their Interactions ................................................................................. 195
• Gene interaction .................................................................................................. 197
• Genes and Chromosomes ..................................................................................... 199
• Eukaryotic Chromosomes .................................................................................... 200
• Gene expression.................................................................................................. 201
• The Genetic code ................................................................................................ 203
• Translation: RNA-Directed Polypeptide Synthesis ................................................... 205
• Regulation of gene expression .............................................................................. 206
• Sex determination and sex linked inheritence .......................................................... 209
• Mutations: Heritable changes in Genes ................................................................... 212
• Transposition...................................................................................................... 216
• Genetic disorders ................................................................................................ 220
• Practice Test Paper-I ........................................................................................... 223
• Practice Test Paper-II .......................................................................................... 237
Chapter 5 Evolutionary Biology.............................................................................. 249–308
• Origin of Earth and Pre-biotic environment ............................................................ 249
• Major episodes in the history of life....................................................................... 250
• Prebiotic Chemical Evolution and the Origion of life................................................ 251
• Oxygenation of Earth........................................................................................... 255
• Geological time scale ........................................................................................... 256
• The History of Evolutionary ................................................................................. 260
• Evidences of evolution ......................................................................................... 261
• Theories of Evolution .......................................................................................... 265
• Population genetics .............................................................................................. 270
• Species Concept and Mechanisms of speciation ..................................................... 272
• Polymorphism .................................................................................................... 278
• Molecular Evolution............................................................................................. 280
• Practice Test Paper-I ........................................................................................... 287
• Practice Test Paper-II .......................................................................................... 298
x CSIR-NET Life Sciences
Chapter 6 Environmental Biology .......................................................................... 309–374
• Major Components of Ecology ............................................................................. 309
• Major Areas of Ecological Study............................................................................ 311
• Population Properties ........................................................................................... 314
– Demographics ........................................................................................ 315
– Survivivorship Curve ............................................................................... 317
– Life Histories .......................................................................................... 319
– Population Growth .................................................................................. 321
– Population Limitating Factors ................................................................... 325
• Community......................................................................................................... 327
– Community Interactions .......................................................................... 327
– Predation ............................................................................................... 327
– Competition between species ................................................................... 331
– Trophic Structure ................................................................................... 333
– Ecological Succession ............................................................................. 334
• Island Biogeography ............................................................................................ 336
• Flow of Energy through Ecosystems .................................................................... 336
– Ecosystems ............................................................................................ 336
– Energy flow ........................................................................................... 337
– Productivity ........................................................................................... 338
– Chemical cycles ...................................................................................... 340
• Behavioural Ecology ............................................................................................ 342
– Finding Food .......................................................................................... 344
– Fighting ................................................................................................. 345
– Finding Sex ............................................................................................ 346
– Cooperation............................................................................................ 348
• Practice Test Paper-I ........................................................................................... 351
• Practice Test Paper-II .......................................................................................... 363
Chapter 7 Biodiversity and Taxonomy .................................................................... 375–420
• Biodiversity ........................................................................................................ 375
– Major biodiversity threats ........................................................................ 376
– Status of Biodiversity in India ................................................................... 377
– Biodiversity hot spots .............................................................................. 380
– Conservation strategies ............................................................................ 382
• DNA finger printing............................................................................................. 385
• Systematics ........................................................................................................ 387
– Classification .......................................................................................... 387
– Phylogeny .............................................................................................. 390
Contents xi
– Biological nomenclature ........................................................................... 391
– Publishing scientific names ...................................................................... 394
• Practice Test Paper-I ........................................................................................... 398
• Practice Test Paper-II .......................................................................................... 409
Model Test papers (1-10) ........................................................................................ 421–548
Seven CSIR NET Life Science Papers (Memory Based) .......................................... 549–632
(Dec., 03 – Dec., 06)
Answer Key ......................................................................................................... 633–656
– Chapter (1–7) .................................................................................. 633–639
– Model Test-papers (1–10) ................................................................. 640–649
– CSIR UGC NET .............................................................................. 650–656
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1
Cell Biology
Cell Biology: St r uct ur e and funct ion of cells and int r acellular or ganelles (of bot h
pr okar yot es and eukar yot es): mechanism of cell division including (mit osis and meiosis)
and cell differ ent iat ion: Cell-cell int er act ion; Malignant gr owt h; Immune r esponse: Dosage
compensat ion and mechanism of sex det er minat ion.
GENERAL INTRODUCTION
1. Re s ol vi n g P owe r : Abilit y t o dist inguish t wo close point s as t wo separ at e point s by
any opt ical syst em is called as it s r esolving power . The r esolving power of human eye
is 100 micr on. Mat hemat ically, r esolving power = l2 wher e l is t h e sou r ce of
illuminat ion. Resolving power of compound micr oscope & elect r on micr oscope is 0.3
micr ons & 10 Å r espect ively.
2. Za ch a r i s J a n s s e n s combined lenses in an effor t t o impr ove magnifying efficiency
and r esolving power . He pr oduced t he fir st compound micr oscope which combines
t wo lenses for gr eat er magnificat ion.
3. Ab ou t Ce l l Con ce p t :
(a) Ma r ce l l o Ma l p i gh i (1628-1694), an It alian micr oscopist , st udied t he st r uct ur e
of plant s. He believed t hat t he plant s ar e composed of separ at e st r uct ur al unit s
which he called “u t r i cl e s ”.
(b) Rob e r t Hook e (1635-1703), examined t hin slices of cor k (dead out er bar k of an
oak) under his micr oscope. He saw hundr eds of ver y small hexagonal ‘boxes’ or
‘chamber s’ which ar e t oget her appear ed like a ‘honeycomb’. Th e t e r m ‘ce l l ’
wa s coi n e d b y Rob e r t Hook e t o denot e t hese chamber s. His obser vat ions,
alongwit h t he figur es, wer e published in 1665 in Micrographia.
(c) An t on von Leeu wen h oek , 1674, using good qualit y simple lenses (magnifying
u pt o 200 t i mes ) obs er ved u n i cel l u l a r or ga n i s ms a n d ca l l ed t h em ‘wi l d
animalcules’. I n t his wa y, he wa s t he fir st t o obser ve “living a nd moving
individual” cells as compar ed t o t he “fixed” cells seen by ear lier wor ker s.
(d) H.J .Du t r och e t (1824), a Fr ench scient ist , boiled some t issues and separ at ed
t he cells fr om one anot her . He expr essed t he idea of individual cells i.e., cells
2 CSIR-NET Life Sciences
wer e not just spaces bet ween a net wor k of fibr es, but t hat t hese wer e separ at e
and separ able unit s.
4. Con ce p t of P r ot op l a s m :
Cor t i (1772) fir st of all obser ved t hat all cells cont ain a living subst ance. This was
fir st obser ved by Cor t i (1772). F e l i x Du j a r d i n (1836), obser ved it in living amoebae,
and called it ‘S arcode’. In 1839, J .E. P u r k i n j e used t he wor d ‘prot oplasm’ t o descr ibe
t he living subst ance. Hugo von Mohl (1846), also suggest ed t he same name – pr ot oplasm
– for t he similar subst ance found in plant cells.
5. Rob e r t Br own , an English nat ur alist , descr ibed in 1828, char act er ist ic dancing of
cell par t icles. It is now, t her efor e, known as Br own i a n move me n t .
6. Nu cle u s : In 1831, Rob e r t Br own saw t hat small spher ical body was pr esent in
ever y plant cell. He used t he wor d ‘nucleus’ t o ident ify t hem.
7. Cell Th eor y : Two Ger man biologist s, M.J . Sch lei d en (1838) and Th eod or Sch wa n n
(1839) pr oposed cell t heor y (or cell doct r ine) – which unified t he ideas pr evailing at
t hat t ime. He st at ed t hat –
(a) living t hings ar e composed of cells and cell pr oduct s.
(b) cells ar e t he fundament al st r uct ur al unit s of living or ganisms.
In fact , Schwann coined t he wor d “me t a b ol i s m” for all chemical pr ocesses car r ied on
in t he cell. Act ually, he called cells “t h e u n i t of li fe ”.
8. Ru d ol f Vi r ch ow (1858), a Ger man pat hologist , developed t he idea of gener at ion t o
gener at ion cont inuit y of cell t hat Omnis cellula e cellula (i.e. cells ar ise fr om pr e-
exist ing cells). This occur s by t he division of cell.
9. Nu c l e o l u s: I n 1781, F on t a n a ha d seen dense spher ica l body inside nucleus.
Sch lei d en (1838) also descr ibed it . But , it was given t he name “nucleolus” by Bowma n
(1840).
10. Na ge li and Cr a mer (1855) gave t he name “cell membr ane” t o t he out er boundar y of
t he pr ot oplasm. Ove r t on (1899) pr oved it s exist ence. J .Q. P l owe (1931), lat er on,
called it ‘plasmalemma’.
11. P r ot op l a s m Th e or y : Ma x Sc h u l t ze (1861) pr oposed t he pr ot opla sm t heor y.
Accor ding t o it “cell is an accumulation of living substance (or protoplasm) which is
limit ed by an out er membrane, and possesses a nucleus”.
12. P l a s t i d s : N. Pr ingsheim and J . Sachs (1865-1892) descr ibed colour ed bodies in t he
cyt oplasm which wer e called plast id by Ha eckel in 1866.
13. Sch i mp e r (1883) classified plast ids int o t hr ee t ypes – Le u cop l a s t s (colour less),
Ch l or op l a s t s (gr een) and Ch r omop l a s t s (colour ot her t han gr een). The gr een
plast ids wer e also ident ified by Meyer (1883) who called t hem aut oplast s. However ,
Er r er a gave t he name chloroplast s t o t hem.
14. Mi t och on d r i a : Kolli ck e r (1880) was t he fir st t o obser ve small t hr ead-like st r uct ur es
in t he cyt oplasm of t he st r iat ed muscle cells of insect . These wer e called ‘fila’ by
F l e mmi n g (1882). Alt ma n n (1890) descr ibed t hem as “bioplast s”. It was Ben d a (1897)
who coined t he t er m mit ochondria.
15. Ce n t r os ome : Bove r i (1888) used t he wor d “cent r osome” for a body found at one
pole of t he cell near t he nucleus in animal cells. Most of t he plant cells wer e found t o
be lacking it .
Cell Biology 3
16. Gol gi a p p a r a t u s : Ca me l l o Gol gi (1898), an It alian scient ist , discover ed in t he
cyt oplasm of ner ve cells of owl/cat , a complex st r uct ur e which he called ‘I n t e r n a l
r e t i cu l a r a p p a r a t u s ’.
17. Pr ot oplasm is a polyphasic cr yst allo-colloidal solut ion. Va r i ou s t h e or i e s a b ou t t h e
n a t u r e of p r ot op l a s m a r e :
(a) Alveolar t heor y of Bu t s c h l i ;
(b) Fibrillar t heor y of Ve lt on ;
(c) Granular t heor y of Al t ma n ;
(d) Colloidal t heor y of Fisch er ;
(e) Reticular t heor y of F r oma n n , and
(f) sol gel t heor y of Hyma n .
But colloidal t heor y of Fischer is best . Conver sion of sol int o gel and vice ver sa is due
t o colloidal nat ur e of cyt oplasm.
18. Cyclosis of cyt opla sm in euka r yot ic cells is due t o sol gel conver sion a nd
micr ofilament act ivit ies.
19. In Paramoecium, cyclosis moves food vacuoles in ‘8’ like manner .
20. Cyt oplasm coagulat es at t emper at ur e above 60°.
21. Amount of wat er in cell is usually not mor e t han 3 quar t er s, i.e. 75%.
22. pH of cyt oplasm, nucleoplasm and human blood is 6.9 ± 0.2, 7.4 ± 0.2 and 7.34 ± 0.2
r espect ively.
23. Pr ot eins and enzymes in t he cyt oplasm ar e found in colloidal for m. This incr eases
t heir sur face ar ea. Vit amins, amino acids, miner als, sugar s and nucleic acids ar e
found in solut ion for m.
24. Cell coat (Gl ycoca l yx or ext r aneous coat ) is made up of oligosacchar ides which act
as r ecognit ion-cent r e dur ing or gan t r ansplant at ion.
25. Swa mme r d a m was fir st t o descr ibe (RBC of fr og). Du t r och e t (1824) gave t he idea
of individualit y of cells.
Ter m cell (L. cella = hollow space) coined by Hook e (1665) is misnomer as cell is not
a hollow st r uct ur e. It has cyt oplasm and cont ains or ganelles, inclusions and nucleus.
Leeu wen h oek (1672) was fir st t o see a fr ee cell under micr oscope and called t hem
t iny animalcules. Ma l p h i gi (1661) called cells as s a ccu le s (ut r icles).
26. Con t r i b u t i on of s ci e n t i s t s i n t h e fi e l d of t i s s u e cu l t u r e
(a) Ha b e r l a n d t (1902) suggest ed t he idea of t issue cult ur e.
(b) St e wa r d et al (1957) pr ovided fir st evidence of cellular t ot ipot ency by gr owing
mat ur e pr oblem t issue of car r ot r oot s in a medium supplement ed wit h coconut
milk.
(c) Tissue cult ur e was r aised by Wh i t e (1932) when he gr ew t omat o r oot s on ar t ificial
medium.
(d) Ca l l u s by Wh i t e , Ga u t h r e t and Nob e cou r t .
(e) Di ffe r e n t i a t i on of Ca l l u s int o t issues by Sk oog a n d Mi l l e r .
(f) S i n gl e cel l cu l t u r e (cellular t ot ipot ency) by St e wa r d (1957).
(g) Nu r s e t e ch n i q u e t o get callus fr om a single cell by Mu i r e t a l (1958).
(h) Mi cr och a m ber t ech n i qu e for single cell cult ur e by Va s i l & Hi l d e r b r a n d t
(1965).
4 CSIR-NET Life Sciences
(i) Em br yoi d (non zygot ic embr yo) cult ur e by St e wa r d e t a l (1963), Ha lp e r i n &
We t h e r e l l (1964).
(j) Em br yo cu l t u r e by La i ba ch (1928).
(k) Pol l en /Ha p l oi d a n d r ogen i c cu l t u r e fr om a nt her s of Da t ur e by Gu h a &
Ma h e s h wa r i (1966).
27. Unicellular eukar yot e is 1-1000 mµ in size.
28. Ost r ich egg (Lar gest cell) is 15-20 × 13.5-15 cm in size.
29. Human ner ve cell (Longest animal cell) is 90 cm.
30. Lar gest acellular plant Acetabularia is 10 cm long.
31. Vir uses do not have a cellular st r uct ur e. Ost r ich egg is not consider ed as t r ue cell as
it st or es a lar ge amount of r eser ve food.
32. In human beings, cells of k i d n e y a r e s ma l l e s t and of ner ve fibr e longest .
33. Sma l l e s t c e l l (Mycoplasma gallisepticum – PPLO) is 0.1 t o 0.3 t o mµ in size.
CELL MEMBRANE
1. All cells ar e enclosed by a t hin, film-like membr ane called t he p l a s ma me mb r a n e
or p l a s ma l e mma .
2. Da n i e l l i a n d Da vs on (1935) pr oposed a “Tr i l a me l l a r mod e l ”. Accor ding t o t his,
t he plasma membr ane is for med of a bimolecular layer of phospholipids (35 Å t hick)
sandwit ched bet ween t wo layer s of pr ot eins (each 20 Å t hick). Thus, t he t ot al t hickness
of plasma membr ane, as per t heir model, should be 20 Å + 35 Å + 20 Å = 75 Å (i.e.,
about 75 Å).
The model was pr oposed even befor e t he plasma membr ane was seen under t he
elect r on micr oscope.
3. J .D. Rob e r t s on (1959) pr oposed a “u n i t me mb r a n e con ce p t ”. Accor ding t o t his, all
biological membr anes shar ed t he same basic st r uct ur e :
(a) These ar e about 75 Å t hick.
(b) These have a char act er ist ics t r ilaminar appear ance when viewed wit h elect r on
micr oscope.
(c) The t hr ee layer s ar e a r esult of t he same ar r angement of pr ot eins and lipids as
pr oposed by Danielli and Davson.
4. Si n ge r a n d Ni col s on (1972) put for war d t he “fl u i d mos a i c mod e l ” of membr ane
st r uct ur e. It is t he lat est and most widely accept ed model. Accor ding t o t his model,
t he cell membr ane consist s of a highly viscous fluid mat r ix of t wo layer s of phospholipids
molecules. These ser ve as a r elat ively imper meable bar r ier t o t he passage of most
wat er soluble molecules. Pr ot ein molecules on t heir complexes occur in t he membr ane,
but not in cont inuous layer ; inst ead, t hese occur as separ at e par t icles asymmet r ically
ar r anged in a mosaic pat t er n. Some of t hese (per ipher al or e xt r i n s i c p r ot e i n s ) ar e
loosely bound at t he polar sur faces of lipid layer s. Ot her s (called int egr al or i n t r i n s i c
p r ot e i n s ), penet r at e deeply int o t he lipid layer . Some of t he int egr al pr ot eins penet r at e
t hr ough t he phospholipids layer s and pr oject on bot h t he sur faces. These ar e called
Tr a n s me mb r a n e s or t u n n e l p r ot e i n s .
5. The plasma membr ane cont ains lipids (32%), pr ot eins (42%), car bohydr at es (6%) and
wat er (20%) alt hough var iat ions ar e always t her e.
Cell Biology 5
6. The car bohydr at es occur only at t he out er sur face of t he membr ane. Their molecules
a r e cova l en t l y l i n ked t o (i) t he pola r hea ds of some lipid molecules (for ming
g l y c o l i p i d s) a n d (ii ) most of t h e pr ot ein s exposed a t ou t er su r fa ce (for min g
gl ycop r ot e i n s ). The car bohydr at es so bound t o membr ane component s const it ut e
t he gl ycoca l yx of cell sur face.
7. Th e s u g a r p o r t i o n s o f g l y c o l i p i d s a n d g l y c o p r o t e i n s a r e i n v o l v e d i n
r e cogn i t i on me ch a n i s ms :-
(a) Sugar r ecognit ion sit es of t wo neighbour ing cells may bind each ot her causing
cell-t o-cell adhesion. This enables cells t o or ient t hemselves and t o for m t issues.
(b) Thr ough glycopr ot eins, ba ct er ia r ecognise ea ch ot her (fema le ba ct er ia a r e
r ecognized by male bact er ia; Paramoecia of differ ent mat ing t ypes r ecognize
each ot her ).
(c) These pr ovide t he basic of immune r esponse and var ious cont r ol syst ems, wher e
glycopr ot eins act as ant igens.
8. Lipids and int egr al pr ot eins ar e a mp h i p a t h i c in nat ur e (i.e. have bot h hydr ophobic
and hydr ophilic gr oups). The hydr ophobic ends ar e sit uat ed inside t he bilayer while
t he hydr ophilic gr oups ar e dir ect ed out war ds. Thus, t he membr ane is held t oget her
pr imar ily by hydr ophobic at t r act ion.
However , t he lipids have links in t heir fat t y acid t ails. These links pr event close
packing of molecules and make t he membr ane st r uct ur e mor e fluid. The fluidit y
incr eases wit h decr easing lengt h of fat t y acid t ails.
9. Thousand of differ ent t ypes of pr ot eins can occur in cell membr anes. These may be
pur ely st r uct ur al (pr ovide elast icit y and mechanical suppor t ) or have addit ional
funct ions as :
(a) Car r ier s, for t r anspor t ing specific molecules int o or out of t he cell.
(b) Recept or s, for immediat e flow of infor mat ion int o t he cells.
10. Gases like O
2
and CO
2
diffuse r apidly in solut ion t hr ough membr anes.
11. Ions and small polar molecules diffuse slowly t hr ough t he membr anes. Unchanged
and fat soluble molecules pass t hr ough membr anes much mor e r apidly.
12. En d oc yt os i s occur s by an infolding or ext ension of plasma membr ane t o for m a
vacuole or a vesicle (small vacuole). It is of t wo t ypes :
(a) P h a gocyt os i s

(i.e. cell eat ing): Mat er ial is t aken up in solid for m. The cells
involved in phagocyt osis ar e called p h a gocyt e s or p h a gocyt i c cells (e.g. whit e
blood cells). The vesicle/vacuole for med is called p h a gocyt i c va cu ol e.
(b) P i n ocyt os i s (i.e. cell dr inking): Mat er ial is t aken up in liquid (solut ion/colloid/
suspension) for m. If t he vesicle for med is ext r emely small, t he pr ocess is known
as mi cr op i n ocyt os i s and t he vesicle is mi cr op i n ocyt ot i c.
13. Exoc yt os i s or r ever se of endocyt osis by which mat er ials ar e r emoved fr om t he cells
including r ever se pinocyt osis.
14. Memb r a n e Ch a n n els: They ar e of t wo t ype’s aqueous channels for t he passage of
wat er and ion channels for t he passage of ions. Nehar and Sakmann got Noble pr ize
for discover y of single ion channels.
15. P s e u d op od i a l Move me n t : Pseudopodia ar e blunt out gr owt hs which ar e for med by
t hr ee development s. Sol-gel changes, cyt oplasmic st r eaming and ext ension of plasma
membr a nce e.g., Amoeba, Macrophages, WBC, et c.
6 CSIR-NET Life Sciences
17. Un d u l a t i on : They ar e small pr ot r usions, pr oject ions or r uffings of t he membr ane
which pass out like a r egular wave in t he ar ea of cont act wit h a solid subst r at um.
STRUCTURAL ORGANISATION OF A CELL
1. Smaller cells wit h smaller volume have mor e sur face ar ea. Su r fa ce : vol u me r a t i o
d e cr e a s e s wi t h i n cr e a s i n g s i ze of ce ll. Lar ger cells incr ease t heir sur face ar ea by
developing a cylindr ical shape or by for ming numer ous ext ensions of t he cell membr ane
like micr ovilli, ER, et c.
2. Smaller cells have mor e sur faces: volume r at io and higher nucleo-cyt oplasmic r at io
hence a r e mor e a ct ive.
3. Posit ion, cell wall, age, viscosit y of cyt oplasm, skelet on and funct ion of t he cell, cont r ol
t he shape of cell, e.g., RBC is biconcave t o incr ease sur face ar ea. Ner ve cells ar e
lar ge as t hey ar e able t o conduct impulses.
4. Cells r egulat e t heir act ivit ies by flow of ener gy and flow of ext r insic and int r insic
(genet ic) infor mat ion.
5. Gr een cells t r ap r adiant solar ener gy and conver t it int o chemical (pot ent ial) ener gy
like ATP which on oxidat ion of food is conver t ed int o kinet ic ener gy for doing wor k.
6. Sch wa n n (1839) r ecognized t hat animal and plant cells ar e alike except t hat animal
cells lack cell wall. Schleiden st at ed t hat cell is t he unit of st r uct ur e and budded off
fr om nucleus. Ru d ol f vi r c h ow (1855) was fir st t o modify cell t heor y and gave
gener alizat ion- “Omnis cellula e cellula”. Vir uses ar e except ion t o cell t heor y. The
specialized cells lose some of t heir aut onomous act ivit ies, e.g., muscle and ner ve
cells do not divide and RBCs do not r espir e.
7. Cells show 3 t ypes of or ganizat ion:
(a) P r ok a r yot i c ce l l s e.g., Bact er ia, cyanobact er ia, ar chaebact er ia, mycoplasma
(PPLO), r icket t siae. Size 0.1 t o 5 µ; DNA : RNA r at io 1 : 2, r -RNA-65%, A + T / G
+ C r at io = 0.88; only one envelope syst em; membr ane bound or ganelles absent ;
hist one, nuclear membr ane, nucleolus, cyclosis meiosis absent .
(b) Eu k a r yot i c ce l l . Size 3 t o 30 µ, DNA : RNA is 1:1, r RNA = 45%, A + T / G + C
r at io = 1.52; t wo envelop syst em, membr ane bound or ganelles; hist one and t r ue
nucleus pr esent .
(c) Me s ok a r yot i c ce ll e.g., Dinoflagellat es (a t ype of algae) – hist one pr ot ein absent
but nucleus pr esent .
8. Cell or ganelles (or ganoids) ar e of four t ypes on t he basis of membr anes.
(i) Or g a n e l l e b o u n d e d b y s i n g l e u n i t m e m b r a n e. e . g. , Mi cr ob od i e s
(per oxisomes, sphaer osomes, glyoxysomes, lysosomes), ER, golgi bodies.
(ii) Or ga n elle bou n d ed by d ou ble membr a n e e.g., Plast ids, mit ochondr ia and
nucleus.
(iii) Or ga n e lle b ou n d e d b y t r i p le me mb r a n e e.g. Tr a nsosomes.
(iv) Or ga n e l l e wi t h ou t a n y me mb r a n e e.g., Ribosomes, cent r iole, nucleolus.
9. Pr ot oplasm of eukar yot ic cells shows of st r eaming movement s known as c yc l os i s :
It is of t wo t ypes.
(i) Rot a t i on (cyt oplasm moves ar ound a vacuole in one dir ect ion) e.g., Hydrilla
leaf cells.
Cell Biology 7
(ii) Ci r cu l a t i on (movement in differ ent dir ect ion ar ound differ ent vacuoles e.g.,
st aminal hair s of Tr adescant ia (Rhoeo discolor ). These movement s ar e due t o
colloidal nat ur e of cyt oplasm and micr ofilament act ivit ies.
10. Ri b os ome s :
(a) Ribosomes wer e fir st discover ed by P a l a d e in animal cell and called t hem as
mi cr os ome s . Rob i n s on a n d Br own discover ed t hem fir st in plant cell. Cla u d e
(1955) called t hese st r uct ur es as r i b os ome s .
(b) Pr ot eins synt hesized on fr ee r ibosome ar e used wit hin cell. Pr ot eins synt hesized
on bound r ibosomes ar e used out side t he cell or incor por at ed int o membr ane or
go out as secr et or y (expor t ) pr ot ein.
(c) They ar e smallest , membr aneless or ganelle and ar e called r i b o-n u cle op r ot e i n
or P a la d e p a r t i cle s . These a r e n e ga t i ve ly ch a r ge d and cont ained r RNA and
pr ot ein and wer e seen only aft er t he discover y of elect r on micr oscope. Their
size is 150 t o 250 A. These act as sit e of pr ot ein synt hesis. A r ibosome may be 70
S or 80 S. (S=sediment at ion coefficient ) and consist s of a smaller and lar ger sub
unit . 70 S ar e found in bot h pr okar yot ic and eukar yot ic cells & lie fr eely in
cyt oplasm as in pr okar yot es. 80 S r ibosomes ar e eit her at t ached t o endoplasmic
r et iculum or nuclea r membr a ne or lie fr eely in cyt opla sm. Er ga s ome s or
p ol yr i b os ome s (Ri ch . 1963) ar e for med by t he combinat ion of 5 t o 6 r ibosomes
on a single mRNA. A 70 S r ibosome has t hr ee molecules of r RNA (16 S, 5 S, 23
S) and 53 pr ot ein molecules and 80 S r ibosome has 3 molecules of r RNA (18 S, 5
S, and 28 S) and 80 molecules of pr ot ein.
11. En d op l a s mi c r e t i cu l u m (Er gast oplasm) : It was r epor t ed by P or t e r , Cla u d e and
F u l l ma n (1945). It was named as endoplasmic r et iculum by Por t er (1953). In muscles,
it is called s a r cop l a s mi c r e t i cu l u m, in eyes called mye loi d b od i e s and in ner ves
as Ni s s le gr a n u le s . ER for ms int r acellular t r anspor t syst em and pr ovides mechanical
suppor t t o cyt oplasm. GERL (Golgi associat ed wit h ER fr om which lysosomes ar ise)
syst em is for med by ER and golgi bodies and for m lysosomes. ER is of t wo t ypes –
Smoot h endoplasmic r et iculum (SER) and Rough endoplasmic r et iculum (RER). RER
a r ises fr om n u c l e a r me mb r a n e. RER is mainly c i s t e r n a l a nd st udded wit h
r ibosomes. SER consist s of t ubules mainly. It const it ut ues mor e t han half of t he t ot al
cell membr anes in a cell. SER helps t o synt hesize lipids and helps in det oxificat ion.
12. P l a s t i d s : They ar e double walled DNA cont aining l a r ge s t or ga n e l l e in plant cells,
discover ed by Ha eckel (1865). These ar e developed fr om colour less pr oplast ids found
in mer ist ems. Thr ee t ypes of plast ids ar e (i) Le u cop l a s t s: Lar gest , colour less, found
in unexposed par t s and st or e st ar ch (a myl op l a s t ), fa t (c l a i op l a s t ) or Pr ot ein
(a l e u r on p l a s t ). (ii) Ch r omop l a s t s : Second lar gest plast ids, have car ont enoids t o
pr ovide at t r act ive colour t o fr uit s, seeds, flower s. (iii) Ch l or op l a s t s : Gr een plast ids
discover ed by Sa ch s (1862) but named Ch l or op l a s t by Sch i mp e r (1885) st or e st ar ch
t empor ar ily; shape var iable, maximum var iat ion in shape is found in gr een algae.
Shape is planoconvex or discoid; each chlor oplast has t wo membr anes. It s mat r ix
(st r oma ) ha s pr oka r yot ic na ked cir cula r DNA (0.5%), RNA, vit a min E a nd K,
plast oglobules (osmiophillic globules), st ar ch par t icles; 70 S r ibosomes, miner als (Fe,
Mg, Cu, Mn, Zn, Co) and enzymes of dar k r eact ion of phot osynt hesis. 50% of mat r ix is
filled wit h Ru b i s co e n zyme. In mat r ix ar e found double membr ane bounded t ubular
sacs called t h yl a k oi d s (s t r u ct u r a l u n i t of ch l or op l a s t ) which ar e st acked t o for m
gr ana; number of gr ana per chlor oplast is 40-100 and each gr anum has 2-100 st acked
8 CSIR-NET Life Sciences
t hylakoids. Inner membr ane of t hylakoid bear s q u a n t a s ome s (funct ional unit of
chlor oplast , discover ed by P a r k a n d Bi ggi n s (1962), size 180 × 150 × 100 Å, called
p h ot os yn t h e t i c u n i t s (PSU) wher e pr imar y act of phot osynt hesis (i.e. r elease of
e

) occur s. A quant asome has 230 chlor ophyll molecules (160 chl a + 70 chl b) and
about 50 car ot enoid molecules. One of t he molecules of Chl a act s as r e a ct i on (t r a p )
ce n t r e. It is P
700
in PS I and P
680
in PS II. Two gr ana ar e joined by F r et s ch a n n el
(st r oma la mella ). Chlor opla st s a nd mit ochondr ia a r e ener gy t r a nsducing. DNA
cont aining, semiaut onomous, double walled or ganelles and called cell wit hin cell
because t hey have t heir own pr ot ein machiner y and show cyt oplasmic inher it ance.
No life is possible on t his ear t h wit hout chlor oplast .
13. Chl a is C
55
H
72
O
5
N
4
Mg – blue gr een and has – CH
3
gr oup.
14. Chl b is C
55
H
70
O
6
N
4
Mg – gr eenish colour and has – CHO gr oup.
15. Pyr enoid is a pr ot einaceous body ar ound which st ar ch is st or ed in gr een algae.
16. Ch l or opl a s t s a r e ext r emel y fr a gi l e os mot i ca l l y a n d bu r s t i n H
2
O a n d h en ce,
chlor oplast s ar e isolat ed fr om gr een leaves using sugar solut ion.
17. Golgi b od i e s : Also called as (Lipochondr ia, Idiosome or Dalt on complex):
They ar e mi d d l e ma n of ce l l and discover ed by Ca me l l o Gol gi (1898) in cyt oplasm
of ner ve cell of owl and cat by s i l ve r me t a l l i c i mp r e gn a t i on t e ch n i q u e. They
for m int er nal r et icular appar at us (apparato recticulare interno) and t ake b l a ck s t a i n
wi t h Su d a n I I I being r ich in lipids. Da l t on a n d F e l i x (1954) obser ved t hem under
TEM and confir med t heir exist ence. In plant s, golgi bodies ar e unconnect ed and
sca t t er ed ca lled d i c t y o s o m e s . I n fu n g i , a d i c t y o s o me i s u n i c i s t e r n a l. I n
ver t ebr at es t hese ar e found near t he nucleus. A dict yosome has a st ack of usually 3-
12 cist er nae wit h swollen ends, t ubules and vesicles. It shows polar it y. Con ca ve or
ma t u r i n g (M) fa ce or t r a n s fa ce is near cell membr ane and ci s or con vex or
for mi n g (F ) fa ce is t owar ds nuclear membr ane. Lysosomes and secr et or y vesicle
ar ise fr om ‘M’ face. New cist er nae ar e for med fr om SER.
Root cap cells ar e r ich in golgi bodies which secr et e mucilage for lubr icat ion of r oot
t ip. They pr ocess package and help in t r anspor t and r elease of secr et or y pr ot eins.
They also cause gl yc os i d a t i on of lipids and gl yc os yl a t i on of pr ot eins t o for m
glycolipids and glycopr ot eins. Golgi body for ms acr osome in sper m, yolk and cor t ical
gr anules in eggs, secr et ion of insulin, lact opr ot ein in mammar y glands, cellulose,
hemicellulose, mucilage, pect in, cell plat e dur ing cell division, r oot hair s et c. They
r egulat e fluid balance of cell. All secr et or y cells ar e r ich in golgi bodies. Ma i n e n zyme
i n gol gi b od i e s i s n u cl e os i d e d i s p h os p h a t a s e. These bodies ar ise fr om SER
mainly.
18. Mi t och on d r i a : They ar e also called as c h on d r i os ome , s a r c os ome , b i op l a s t ,
p l a s t och on d r i a :
They ar e power house of cell, l a r ge s t or ga n e l l e in animal cell and 2
n d
l a r ge s t
or ga n e l l e in plant cell. These ar e double walled, DNA cont aining, self r eplicat ing,
semiaut onomous, or ganelle found only in eukar yot ic aer obic cells (except mat ur e
RBC), fir st obser ved in insect st r iat ed muscles as gr anular st r uct ur e by Kolliker
(1850). He called t hese gr anules of st r iat ed muscles as sar cosomes. Al t ma n (1890)
st udied t hem in det ail and called t hem as b i op l a s t s . He consider ed t hem as symbiont s
compar able t o bact er ia. F l e mmi n g called t hem file and t er m ‘mi t och on d r i a ’ was
Cell Biology 9
used by Ben d a (1897) who st ained t hem wit h J a n u s gr e e n B (a vit al st ain); size
1 = 10µ × 0.2 – 1 µ; Number 1 per cell in Microst erias and Trypanosoma; 50,000 in
Choas chaos and 30,000 t o 3 lakhs in oocyt es of sea ur chin.
A mit ochondr ion has t wo chamber s and t wo membr anes. Inner membr ane is folded
to form cr i s t a e which bear oxysomes (F
0
-F
1
par t icles, element ar y par t icles, ATPase
particles. (Fernandez and M oran particles). Oxys ome s ar e 10
4
t o 10
5
in number ,
called funct ional unit of mit ochondr ia, discover ed by Fer n a n d ez-Mor a n (1961).
Inner chamber has a double st r anded, naked cir cular 5 µ long pr okar yot ic DNA wit h
high G-C r at io, 70S r ibosomes, RNA and 70 t ypes of enzymes. 70% of t ot al enzymes of
cell ar e found in mit ochondr ia. This DNA is 1% of t ot al DNA of cell and discover ed by
Nass (1966). Kr eb’s cycle and oxidat ive phosphor ylat ion occur her e. Mit ochondr ia ar e
called cell wi t h i n ce l l .
19. Mi cr ob od i e s : These ar e (lysosomes, sphaer osomes, glyoxysomes and per oxisomes):
They ar e smallest single membr ane bounded or ganelle. Lys os ome s (suicidal bags,
disposal unit s, scavenger of cell) ar e micr obodies of 0.2-0.8 µ size filled wit h 40 t ypes
of acid hydr olases t o digest (aut olysis) almost ever y t ype of or ganic mat t er except
cellulose. They ar e common in WBC, liver , spleen et c. They wor k at pH-5 and cause
lysis of for eign body; discover ed accident ly by Ch r i st i a n d e Du ve (1955) fr om r at
liver . Novi k off (1956) obser ved t hem under TEM. They show p l e omor p h i s m
(p ol ymor p h i s m).
They ar e of four t ypes: (i) P r i ma r y l ys os ome s (St or a ge gr a n u l e s ) have inact ive
enzymes; (ii) Se con d a r y l ys os ome s (He t e r op h a gos ome s or Di ge s t i ve va cu ol e s
or P h a gol ys os ome s ) cause digest ion (He t e r op h a gy); (iii) Te r t i a r y lys os ome s
(Re s i d u a l b od i e s or Te l ol ys os ome s ) r emoves wast es by ephagy fr om cell by act ing
a s sca ven ger or disposa l u n it ; (iv ) Au t op h a gi c va c u ol e s (Cyt ol ys os ome or
Au t op h a gos ome s ) ar e complex lysosomes and digest old / injur ed / dead cells and
t issues t o keep cell healt hy (aut olysis or aut odigest ion). Scavenging, disappear ance
of t ail, soft ening of gums, acr osomal act ivit y of sper m ar e few ot her examples.
Au t op h a gy is digest ion of st or ed food (glycogens, fat and pr ot eins) dur ing st ar vat ion
t o pr ovide ener gy. Thus aut ophagy has no r ole in scavenging. If lysosomes bur st and
r elease t heir enzymes, t he ent ir e cell is digest ed and liquefied. It is called aut olysis.
Ost eoclast s (which digest bones) ar e r ich in lysosomes; P l a n t ce l l s l a ck l ys os ome s
e xce p t Ne u r os p or a , r oot t i p of ma i ze , ye a s t a n d s e e d s of p e a a n d cot t on .
Lack of Lysosome cause Pombe’s Disease. They ar e also involved wit h diseases such
as Tay-Sachs Syndr ome and Rheumat ic Ar t hr it is (An Aut oimmune disease).
Typ es of Mi cr ob od i es
(a ) Soga e r i s i n e s (Plant lysosomes) ar e micr o bodies filled wit h hydr olyt ic enzymes
for fat synt hesis. They ar e highly r efr act ile and r ich in fat (98%) and t ake black
st ain wit h Sudan iii / Osmium t et r aoxide. These ar e abundant in endosper m of
oily seeds.
(b) Tr a n s os ome s ar e t r iple la yer bounded or ganelle in ovar y follicle cells of bir ds
t o help in yolk for mat ion.
(c) Loma s ome s ar e boder bodies bet ween cell wall and cell membr ane, common in
fungi, discover ed by Moor e a n d Mc Al l i s t e r (1961) and help in cell pr olifer at ion
and elongat ion for diffusion of subst ances r equir ed in cell wall for mat ion.
10 CSIR-NET Life Sciences
(d) P e r oxi s ome s (ur icosomes) ar e micr obodies cont aining enzymes for per oxide
for mat ion. Ca t a l a s e a n d p e r oxi d a s e a r e l a r ge s t a n d s ma l l e s t e n zyme s
found in per oxisomes. In plant s t hey do phot o-r espir at ion in C
3
plant s (Tolber t ,
1972). In animals t hey t ake par t in lipid synt hesis (b-oxidat ion of fat t y acids).
(e) Gl yoxys ome s - Lar gest micr obody of size upt o 10 µ and similar t o per oxisomes
as t hese cont ain cat alase and ot her enzymes for glyoxylat e cycle (a modified
Kr ebs’ cycle in which fat s ar e conver t ed int o car bohydr at es). These ar e common
in ger minat ing oil seeds of cast or , gr oundnut and cucumber s and disappear aft er
ger minat ion.
20. Ce n t r i ole s : They ar e minut e submicr oscopic subcylindr ical st r uct ur es of 300-500
nm lengt h and 150 nm diamet er and usually occur in pair s (diplosome) inside a
specialized fibr ous cyt oplasm called cent r ospher e. The complex is called ce n t r os ome
or c e n t r a l a p p a r a t u s. Ea ch cent r iole ha s a whor l of nine t r iplet fibr ils wit h
int er connect ions amongst adjacent t r iplet s (called C-A linker s) as well as wit h a cent r al
hub t hr ough spokes. Thus show 9 ÷ 0 or ganisat ion. Cent r ioles ar e sur r ounded by
ma s s u l e s or nuclea t ing cent r e or per icent r iola r sa t ellit es for for ma t ion of new
cent r ioles. Cent r ioles ar e r equir ed t o for m basal bodies, cilia, flagella and spindle
poles. They occur in most animal cells except some pr ot ozoan pr ot est s like Amoeba,
common in flagellat e for ms (e.g., many gr een algae, br yophyt es, pt er idophyt es and
cycads). Cent r iole is r ich in t u b u l i n , ATP . Cent r ioles a r e ba sica lly locomot or y
st r uct ur es and t heir r ole in cell division t o for m spindle is secondar y.
Ba s a l b od i e s or basal gr anules or blephar oplast s ar e micr ocylinder s t hat lie below
t he plasmalemma at t he base of flagella and cilia. The st r uct ur e is exact ly similar t o
cent r iole.
21. Ci l i a a n d fl a ge l l a : They ar e micr ot ubular vibr at ile pr opoplasmic pr ocesses st udied
by En gl e ma n and have four par t s: basal body, r oot let s, basal plat e and shaft . Shaft
cont ains of an ext er nal membr ane (ext ension of plasmalemma), a semifluid mat r ix
and an a xon e me. Axoneme has nine per ipher al doublet fibr ils and t wo cent r al singlet
fibr ils. Thus show 9 + 2 fibr illar or ganizat ion. All t he per ipher al doublet fibr ils ar e
int er connect ed by C-A linker s of pr ot ein n exi n . Subfibr e A of each doublet has t wo
bent ar ms, t he out er one wit h a hook. The cent r al fibr ils and side ar ms of subfibr e A
ar e made of d ye n i n pr ot ein wit h ATP-ase act ivit y. Cilia ar e shor t er (5-10 µm as
compar ed t o 150 µm for flagella), mor e numer ous, have sweeping or pendular movement
and beat in a coor dinat ed r hyt hmic movement .
22. Va cu oles : Vacuoles in plant s wer e r epor t ed by Sp a lla n za n i . They ar e noncyt oplasmic
fluid filled, lifeless sacs which ar e separ at ed fr om cyt oplasm by a membr ane called
t on op l a s t .
(a) Sa p va cu ole s : They enclose sap or wat er wit h dissolved inor ganic and or ganic
subst ances. A mat ur e plant cell has a single lar ge cent r al vacuole. Animal cells
have numer ous small sap vacuoles t hat maint ain osmot ic pr essur e. Cell sap is
slight ly acidic and cont ains acids, est er s, phenols, or ganic acids (acet ic and for mic
acids), enzymes, t annin cr yst als and pigment s et c.
(b) Con t r a ct i le va cu ole s : They occur in some simple fr esh wat er for ms (e.g.,
Amoeba, Paramecium, Chlamydomonas ). They pick up wat er fr om sur r ounding
cyt oplasm, expand (diast ole) and collapse (syst ole) t o t hr ow wat er out side.
Cont r act ile vacuoles per for m osmor egulat ion and excr et ion.
Cell Biology 11
(c) Ga s va c u ol e s ( = p s e u d o v a c u o l e s ): Ga s or a i r va cu ol es occu r i n s ome
pr okar yot es. Gas vacuoles st or e met abolic gases and t ake par t in buoyancy
r egulat ion.
23. Nu cle u s : A cell may be uni or mult inucleat ed. If a mult inucleat ed condit ion ar ises
due t o fusion of cells, it is called s yn c yt i u m e.g., plasmodium, body of slime moulds,
young xylem vessels and if due t o r epeat ed nuclear divisions wit hout cyt okinesis, it is
called coen ocyt i c e.g. Vaucheria, Rhizopus. Nucleus was r epor t ed by Rob e r t Br own
(1831) in or chid cells. St r a sb u r ger (1882) pr oved t hat nucleus ar ises fr om pr e exist ing
nucleus by division. He r t wi g and va n Ben eden showed t he r ole of nucleus in
fer t ilizat ion. Ha mme r l i n g (1953) by his gr aft ing exper iment s on Acetabularia (lar gest
unicellular gr een, mar ine alga) pr oved t he r ole of nucleus in her edit y, gr owt h and
mor phology. 1/10
t h
of volume of cell is occupied by nucleus. In a cell, t her e is a
definit e nucleo-cyt oplasmic r at io. Nucleocyt oplasmic index is Volume of nucleus/
(Volume of cell – Volume of nucleus).
About 10% of nuclear membr ane bear s simple of compound por es. Nucleus has 80%
pr ot eins (65% n on h i s t on e, Mol. Wt . high, r ich in t yr os i n e and t r yp t op h a n , acidic
and for ms enzymes and helps in RNA t r ansact ion. 15% pr ot eins ar e basic, h i s t on e
pr ot eins, Mol. Wt . low, r ich in l ys i n e and a r gi n i n e. DNA : hist one r at io 1 : 1.
Nu cle os ome s ar e st r uct ur al unit s of chr omat in. Ter m was given by Ou t d et . A
nucleosome is an oct a me r of h i s t on e pr ot eins and has a cor e of 8 molecules of
hist one pr ot eins (t wo each of H
2
A, H
2
B, H
3
, H
4
) bounded by 13/4 t ur ns of DNA having
about 166 base pair s. H
1
hist one does not for m nucleosome. Size of a chr omosome
var ies fr om 0.5 t o 32 µ. Minimum number of chr omosomes n = 2 e.g., Haplopappus.
Maximum number is 2n = 1262 in Adder ’s fer n (Ophioglossuin). In animals, minimum
number is 2n = 2 in Ascaris sp. And maximum number is 2n = 1600 in Aulacantha and
Radiolar ians.
24. Gi a n t Ch r o mo s o me s a r e: (i) Sa liva r y gla nd chr omosomes (size 2000 µ
m
) (ii )
lampbr ush chr omosomes (size 5900 µ
m
). Those chr omosomes help in r apid synt hesis
of pr ot eins.
25. Nu cl e ol u s : It is a sit e of r ibosome synt hesis. Nucleolus was discover ed by F on t a n a
(1781), descr ibed by Wa gn er and numbed by Bowma n . Ther e is at least one nucleolus
per haploid set of chr omosomes in a cell.
26. Ce l l i n cl u s i on s : Er gast ic, deut oplasmic, par aplasmic bodies ar e non living, non
cyt oplasmic subst ances in vacuoles or cell wall or cyt oplasm of eukar yot ic cells also
called met aplast or deut oplast and ar e of t hr ee t ypes:
(a) Re s e r ve food ma t e r i a ls : It can be st ar ch as in plant cells, glycogen as in animal
cells and fungi, fat or aleur one gr ains (pr ot ein r ich, found as out er most layer of
cells or endosper ms or cer eal gr ains). An aleur one gr ain in made up of a lar ge
par t called c r ys t a l l oi d and a small par t called glob oi d . Cr yst alloid cont ains
nit r ogen as amides. St ar ch is found as gr ains; simple or compound, concent r ic
or acent r ic.
(b) Excr e t or y p r od u ct s : These ar e wast e (end) pr oduct s and useless t o plant . They
get accumulat ed in bar k, old leaves, vacuoles and flower s e.g., alkaloids (Quinine,
At r opine, Canada Balsam: a mount ing agent fr om Abies st em), gums, or ganic
acids, cow milk, lat ex fr om Cow t r ee (Brosimum) and miner al cr yst als.
12 CSIR-NET Life Sciences
Mi n e r a l cr ys t a l s a r e:
(i) CaCO
3
cr yst als occur r ing as a mass of cr yst als in a cellulose wall t o for m
bunch of gr apes called c ys t ol i t h , e.g., banyan leaf cell. In Momordica,
cyst olit h is double and in J ust icia it is wor m like.
(ii) Cr yst als of calcium oxalat e ar e ver y common and called r a p h i d e s (needles
like e.g., Lemna, Pistia), st ar like (s p h a e r a p h i d e s or d r u s e e.g. Colocasia,
Nerium), pr ism like in onion scales or cr yst al sand in At r opa. A cell wit h
r aphides is called i d i ob l a s t .
(iii) Silica on mar gin of leaves of gr asses.
(c) Se cr e t or y p r od u ct s : They ar e useful t o plant s e.g., pigment s, nect ar , essent ial
oil, enzymes et c. Essent ial oils ar e et her eal oils and used in per fumer y.
27. Cyt os k e l t a l s t r u ct u r e s : These ar e fibr ous or fine t ubular st r uct ur es which for m
t he suppor t ive st r uct ur es of t he cell. These a r e of t hr ee t ypes – micr ot ubules,
micr ofilament s and Int er mediat e filament s
(a) Mi cr ot u b u le s d i s cove r e d b y Rob e r t i s a n d F r a n ch i , (1953) t er m coined by
Sla u t t er ba ck (1963), ar e unbr anched hollow non cont r act ile t ubules of indefinit e
lengt h, 25 nm in t hickness wit h 15 nm cor e and for med of 13 helically ar r anged
p r ot ofi l a me n t s of a and b-t ubilin pr ot ein. Micr ot ubules gr ow fr om nucleat ing
cent r es. Micr ot ubules ar e basic st r uct ur es of spindle appar at us, cent r ioles, basal
bodies, cilia and flagella and ar e r esponsible for cell mot ilit y and maint enance of
sha pe. Their t ips ca n gr ow a nd shor t en quickly. GTP, Ca 2+, Mg2+ a nd a
calmodulin bound pr ot ein ar e r equir ed for assembly. Colchicine pr event s it .
Micr ot ubules ar e basic st r uct ur es of spindle appar at us, cent r ioles, basal bodies,
cilia and flagella. They ar e also pr esent in ot her cellular st r uct ur es like sensor y
hair , ner ve pr ocesses, sper m t ail, et c. Micr ot ubules pr esent in cyt oplasm pr ovide
shape and polar it y t o cells. Micr ot ubules ar e absent in pr ocar yot es (except
Anabaena),Amoeba and Slime Moulds.
(b) Mi cr ofi l a me n t s ar e cylindr ical solid, cont r act ile r ods or filament s of act in and
myosin pr ot ein wit h a diamet er of 6 t o 10 nm. Micr ofilament s can for m hexagonal
bundles, t ake par t in cyt oplasmic st r eaming, membr ane undulat ions, cleavage,
cont r act ion of muscles, movement of micr ovilli t o absor b food and endocyt osis.
(c) I n t e r me d i a t e F i l a me n t s ar e int er mediat e in size having diamet er ar ound 10-
15 nm and ar e composed of non-cont r act ile pr ot eins. Int er mediat e fibr es (IF)
ar e of four t ypes–ker at in filament s, neur ofibr ils, glial filament s and het er ogeneous
filament s (viz., desmin filament s, viment in filament s, synemin filament s).They
pr ivide r igidit y t o cell and maint ains t he cell st r uct ur e
CELL CYCLE
I. Interphase and the Control of Cell Division
1. Int er phase is t he per iod bet ween divisions of t he cyt oplasm. A t ypical eukar yot ic cell
will spend most of it s life in int er phase. Some cells lose t he capacit y t o divide alt oget her
and st ay in int er phase indefinit ely. Examples of such cells in humans ar e ner ve cells
and muscle cells. Ot her cells divide r egular ly, ot her s occasionally.
2. Most cells have t wo major phases: mit osis and int er phase oft en r efer r ed t o as t he cell
cycle.
Cell Biology 13
3. For most t issues at any given t ime, only a few cells ar e in mit osis, and most ar e in
int er phase.
4. Int er phase consist s of t hr ee sub-phases.
• G1 is Gap 1, t he per iod just aft er mit osis and befor e t he beginning of DNA
synt hesis.
• Next is S (synt hesis), which is t he t ime when t he cell’s DNA is r eplicat ed.
• G2 is t he t ime aft er S and pr ior t o mit osis.
5. Mit osis and cyt okinesis ar e r efer r ed t o as M phase.
6. The G1-t o-S t r ansit ion commit s t he cell t o ent er anot her cell cycle.
II. Cyclins and other proteins signal events in the cell cycle
1. Tr ansit ions fr om G1 t o S and G2 t o M depend on act ivat ion of a pr ot ein called cyclin-
dependent kinase, or Cdk.
2. A kinase is an enzyme t hat t r ansfer s a phosphat e fr om ATP t o differ ent pr ot ein(s).
This is called phosphor ylat ion.
3. Act ivat ed Cdk t r ansfer s phosphat es fr om ATP t o cer t ain amino acids of pr ot eins t hat
t hen move t he cell in t he dir ect ion of cycling.
4. The Cdk effect on t he cell cycle is a common mechanism in eukar yot ic cells.
• St udies in sea ur chin eggs uncover ed a pr ot ein called t he mat ur at ion pr omot ing
fact or .
• A mut ant yeast t hat lacked Cdk was found, which st alled at t he G1–S boundar y.
• These t wo pr ot eins, one fr om sea ur chins and t he ot her fr om yeast , wer e similar
in st r uct ur e and funct ion. Ot her Cdks have been found in ot her or ganisms,
including humans.
5. Cyclin is a pr ot ein t hat int er act s wit h Cdk. Cyclin binding of Cdk exposes t he act ive
sit e of t he kinase.
6. The cyclin-Cdk complex act s as a pr ot ein kinase t hat t r igger s t r ansit ion fr om G1 t o S.
The cyclin t hen br eaks down and t he Cdk becomes inact ive. Sever al differ ent cyclins
exist , which, when bound t o Cdk, phosphor ylat e differ ent t ar get pr ot eins.
• Cyclin D-Cdk4 act s dur ing t he middle of G1. This is t he r est r ict ion point in G1,
beyond which t he r est of t he cell cycle is inevit able.
• Cyclin E-Cdk2 act s at t he boundar y of G1 t o S t o init iat e DNA r eplicat ion.
• Cyclin A-Cdk2 act s dur ing S and also st imulat es DNA r eplicat ion.
• Cyclin B-Cdk1 act s at t he G2-t o-M boundar y, init iat ing mit osis.
7. Cyclin-Cdk complexes act as checkpoint s. When funct ioning pr oper ly, t hey allow or
pr event t he pa ssa ge t o t he next cell cycle st a ge, depending on t he ext r a - a nd
int r acellular condit ions.
• An example is t he effect of p21 on t he G1-t o-S phase t r ansit ion.
• If DNA is damaged by UV r adiat ion, p21 is synt hesized (a pr ot ein of 21,000
dalt ons).
• It binds t o t he t wo differ ent t ypes of G1 Cdk molecules, pr event ing t heir act ivat ion
unt il damaged DNA is r epair ed. The p21 is t hen degr aded, allowing t he cell
cycle t o pr oceed.
14 CSIR-NET Life Sciences
8. Some t ar get s for cyclin-Cdk complexes include pr ot eins t hat condense chr omosomes
and ot her s t hat cause fr agment at ion of t he nuclear envelope.
9. Cyclin-Cdk defect s have been found in some cancer cells.
• A br east cancer wit h t oo much cyclin D has been found.
• The pr ot ein p53, which inhibit s act ivat ion of Cdk, is found defect ive in half of all
human cancer s.
III. Growth factors can stimulate cells to divide
1. Cyclin-Cdk complexes pr ovide int er nal cont r ol for cell cycle decisions.
2. Cells in mult icellular or ganisms must divide only when appr opr iat e. They must
r espond t o ext er nal signals, cont r ols called gr owt h fact or s.
3. Some cells r espond t o gr owt h fact or s pr ovided by ot her cells.
• Plat elet s r elease plat elet -der ived gr owt h fact or , which diffuses t o t he sur face of
cells t o st imulat e wound healing.
• Int er leukins ar e r eleased fr om one t ype of blood cell t o st imulat e division of
anot her t ype r esult ing in body immune syst em defenses.
• The cells of t he kidney make er yt hr opoiet in, which st imulat es bone mar r ow
cells t o divide and differ ent iat e int o r ed blood cells.
4. Cancer cells cycle inappr opr iat ely because t hey eit her make t heir own gr owt h fact or s
or no longer r equir e t hem t o st ar t cycling.
IV. Regulation of the Cell Cycle
1. Cell cycle is dr iven by specific chemical signals in t he cyt oplasm.
2. M phase cells t hat ar e fused wit h any ot her phase cell, t he lat t er cell will ent er
mit osis.
3. Ce ll cycle con t r ol s ys t e m t r igger s and coor dinat es key event s in t he cell cycle.
4. Cell cycle ch e ck p oi n t s act as st op and go signals. Thr ee major checkpoint s found in
G
1
, G
2
, and M p h a s e s .
(a) G
1
is cr it ical checkpoint . If cells make it past G1, t he ent ir e cell cycle is complet ed.
(b) Non-dividing cells ar e in G
0
s t a t e.
5. Fluct uat ions in cell cycle cont r ol molecules abundance and act ivit y cont r ol cell cycle.
• P r ot e i n k i n a se s ar e act ivat ed by cycli n pr ot eins.
• Act ivit y of pr ot ein kinase is cor r elat ed wit h concent r at ion of specific cyclin (cyclin
dependent kinase or “Cd k ”).
6. MP F (mat ur at ion pr omot ing fact or ) was fir st Cdk descr ibed.
• Cyclin level r ises dur ing int er phase.
• At G2, enough act ive MPF (cyclin + Cdk) is pr esent t o pr omot e mit osis.
• Numer ous phosphor ylat ion event s t hat cause nuclear envelope t o fr agment
and act ivat e ot her enzymes.
• Cyclin is br oken down by pr ot eolyt ic cleavage (MPF inact ive) and Cdk is r ecycled.
• Pr ot eolysis also dr ives M-phase past anaphase by br eaking down pr ot eins t hat
hold sist er chr omat ids t oget her .
Cell Biology 15
7. I n t e r n a l a n d Ext e r n a l Cu e s Re gu l a t e Ce l l Cycl e I n t e r n a l s i gn a l delays st ar t of
anaphase (separ at ion of chr omosomes) unt il all kinet ochor es ar e at t ached t o spindle
fiber s.
• Anaphase pr omot ing complex (APC) is kept in inact ive st at e by pr ot eins associat ed
wit h kinet ochor es.
• Signal ceases when all kinet ochor es ar e at t ached.
8. Gr owt h fa ct or s ar e e xt e r n a l s i gn a l s t hat st imulat e cells t o divide.
• Plat elet der ived gr owt h fact or (PDGF) is r equir ed for division of fibr oblast s.
• PDGF binds t o t yr osine kinase r ecept or s on sur face of cells and t r igger s signal
t r ansduct ion pat hway.
9. De n s i t y d e p e n d e n t i n h i b i t i on descr ibes phenomenon wher eby cells st op gr owing
aft er r eaching a cer t ain densit y. Gr owt h is limit ed by availabilit y of gr owt h fact or .
10. Ca n ce r Ce lls Ha ve Es ca p e d Ce ll Cycle Con t r ols
• Cancer cells do not exhibit densit y dependent inhibit ion.
• Cancer cells do not st op gr owing when gr owt h fact or is deplet ed.
• Cancer cells st op at r andom point s in cell cycle (not checkpoint s).
• Some cancer cell lines ar e i mmor t a l and can divide indefinit ely given t he r ight
ingr edient s.e.g. HeLa cells.
• p53 gene mut at ions in t umor suppr essor genes (e.g. p53) r esult in cancer funct ional
p53 aids cell in checkpoint cont r ol at G1 and G2
CELL REPRODUCTION & CELL DIVISION
I. Systems of Cell Reproduction
1. Four event s occur befor e and dur ing cell division.
• A signal t o r epr oduce must be r eceived.
• Replicat ion of DNA and vit al cell component s must occur .
• DNA must dist r ibut e t o t he new cells.
• The cell membr ane (and cell wall in some or ganisms) must separ at e t he t wo
new cells.
II. Prokaryotes divide by fission
1. Pr okar yot ic cells gr ow in size, r eplicat e DNA, and divide int o t wo new cells. This
pr ocess is called fission.
2. Escherichia coli (a bact er ium) simply divides as quickly as r esour ces per mit . At 37
o
C,
t his is about once ever y 40 minut es. When r esour ces ar e abundant , E. coli can divide
ever y 20 minut es.
3. Pr okar yot es gener ally have just one cir cular chr omosome.
• The E. coli chr omosome is 1.6 mm in diamet er , making t he unfolded cir cle 100
t imes gr eat er t han t he size of t he cell. The molecule is packaged by folding in on
it self wit h t he aid of basic pr ot eins t hat associat e wit h t he acidic DNA.
• Cir cular chr omosomes appear t o be char act er ist ic of all pr okar yot es.
16 CSIR-NET Life Sciences
4. The pr okar yot es have a sit e called ori, wher e DNA r eplicat ion begins, and a sit e t er,
wher e it ends.
• Ori is shor t for or igin of r eplicat ion.
• Ter is shor t for t er minus of r eplicat ion.
5. As DNA r eplicat es, each of t he t wo r esult ing DNA molecules at t aches t o t he plasma
membr ane. As t he bact er ium gr ows, new plasma membr ane is added bet ween t he
at t achment point s, and t he DNA molecules ar e moved apar t .
6. Cyt okinesis, which is cell par t it ioning, begins ar ound 20 minut es aft er chr omosome
duplicat ion is complet ed. A pinching of t he plasma membr ane t o for m a const r ict ing
r ing separ at es t he one cell int o t wo, each wit h a complet e chr omosome.
• A t ubulin-like fiber is involved in t he pur se-st r ing const r ict ion.
III. Eukaryotic cells divide by mitosis or meiosis
1. All r epr oduct ion involves r epr oduct ion signals, DNA r eplicat ion, segr egat ion, and
cyt okinesis.
2. Unlike pr okar yot es, eukar yot ic cells do not const ant ly divide whenever envir onment al
condit ions ar e adequat e, alt hough unicellular eukar yot es do so mor e oft en t han t he
cells of mult icellular or ganisms.
• Some differ ent iat ed cells of mult icellular or ganisms r ar ely or never divide.
• Signals t o divide ar e r elat ed t o t he needs of t he ent ir e or ganism, not simply t he
oppor t unit y cr eat ed by r esour ces.
3. Eukar yot es usually have many chr omosomes. Eukar yot es have a nucleus, which
must r eplicat e and, wit h few except ions, divide dur ing cell division. Mit osis gener at es
t wo cells wit h t he same genet ic infor mat ion as t he or iginal cell.
4. Meiosis is a specia lized cell division used for sexua l r epr oduct ion. The genet ic
infor mat ion of t he chr omosomes is shuffled, and t he cells, called gamet es, t ypically
get one-half of t he or iginal DNA complement .
IV. Mitosis: Distributing Exact Copies of Genetic Information
1. A single nucleus gives r ise t o t wo genet ically ident ical nuclei, one for each of t he t wo
new daught er cells.
2. Mit osis is a cont inuous event , but it is convenient t o look at it as a ser ies of st eps.
• When t he cell ent er s S phase and DNA is r eplicat ed, t he cent r osome r eplicat es
t o for m t wo cent r osomes. This event is cont r olled by cyclin E-Cdk2, whose
concent r at ion peaks at t he G1-t o-S t r ansit ion. This is t he key event init iat ing
t he dir ect ion of mit osis.
• Dur ing G2-t o-M t r ansit ion, t he t wo cent r osomes separ at e fr om each ot her and
move t o opposit e ends of t he nuclear envelope. The or ient at ion of t he cent r osomes
det er mines t he cell’s plane of division.
• In many or ganisms, each cent r osome cont ains a pair of cent r ioles t hat have
r eplicat ed dur ing int er phase. Cent r osomes ar e r egions wher e micr ot ubules for m.
These micr ot ubules will or chest r at e t he movement of chr omosomes.
Cell Biology 17
The spindle forms during prophase
3. In pr ophase, polar micr ot ubules for m bet ween t he t wo cent r osomes and make up t he
developing spindle.
4. Each polar micr ot ubule r uns fr om one mit ot ic cent er t o just beyond t he middle of t he
spindle, wher e it over laps and int er act s wit h a micr ot ubule fr om t he ot her side.
Initially, these m icrotubules are constantly form ing and depolymerizing (“falling apart ”)
dur ing t his per iod. Recall t hat micr ot ubules gr ow by addit ion of t ubulin dimer s t o t he
+ en d of t h e mi cr ot u bu l e. Wh en mi cr ot u bu l es fr om on e cen t r os ome con t a ct
micr ot ubules fr om t he ot her , t hey become mor e st able.
5. The mit ot ic spindle ser ves as a “r ailr oad t r ack” along which chr omosomes will move
lat er in mit osis.
A prophase chromosome consists of two chromatids
6. Dur ing pr ophase, chr omosomes compact and coil, becoming mor e dense. Pr ophase
chr omosomes consist of t wo chr omat ids, held t oget her over much of t heir lengt h.
The r egion of t ight binding bet ween t he chr omat ids, t he cent r omer e, is wher e t he
micr ot ubules will associat e wit h t he chr omat ids.
7. Lat e in pr ophase, t he kinet ochor es develop. The kinet ochor e is locat ed in t he r egion
ar ound t he cent r omer e and is t he sit e wher e micr ot ubules at t ach t o t he chr omat ids.
Chromosome movements are highly organized
8. The movement phases of chr omosomes ar e designat ed pr o-met aphase, met aphase,
and anaphase.
9. Dur ing pr o-met aphase, t he nuclear lamina disint egr at es and t he nuclear envelope
br eaks int o small vesicles per mit t ing t he fiber s of t he spindle t o “invade” t he nuclear
r egion.
• The spindle micr ot ubules t hen associat e wit h kinet ochor es.
• These ar e called kinet ochor e micr ot ubules.
• The micr ot ubules fr om one pole associat e wit h t he kinet ochor e of one of t he
member s of a pair of chr omat ids. Micr ot ubules fr om t he ot her pole associat e
wit h t he kinet ochor e of t he ot her member .
• Repulsive for ces fr om t he poles push chr omosomes t owa r d t he cent er , or
equat or ial plat e, in a r at her aimless back and for t h mot ion.
• The t wo chr omat ids ar e held t oget her , pr esumably by pr ot eins called coh e s i n s .
10. Dur ing met aphase, t he kinet ochor es ar r ive at t he equat or ial plat e.
• Chr omosomes ar e fully condensed and have dist inguishable shapes.
• Cohesins br eak down.
• DNA t opoisomer a se I I unr a vels t he int er connect ed DNA molecules a t t he
cent r omer e, and all t he chr omat ids separ at e simult aneously.
11. Anaphase begins when t he cent r omer es separ at e.
• The pr ocess t akes 10 t o 60 minut es for t he chr omosomes t o move t o opposit e
poles.
• Molecular mot or s at t he kinet ochor es move t he chr omosomes t owar d t he poles,
account ing for about 75% of t he mot ion.
• About 25% of t he mot ion comes fr om shor t ening of t he micr ot ubules at t he
poles.
18 CSIR-NET Life Sciences
• Addit ional dist ance is gained by t he separ at ing of t he mit ot ic cent er s. This
incr ease in dist ance bet ween t he poles is done by t he polar micr ot ubules, which
have mot or pr ot eins associat ed in t he over lapping r egions. By t his pr ocess t he
dist ance bet ween t he poles doubles.
Nuclei re-form during telophase
12. When chr omosomes finish moving, t elophase begins. Nuclear envelopes and nucleoli
coalesce and r e-for m.
V. Cytokinesis: The Division of the Cytoplasm
1. Animal cells divide by a fur r owing (a “pinching in” or const r ict ion) of t he plasma
membr a ne.
2. Micr ofilament s of act in and t he mot or pr ot ein filament myosin fir st for m a r ing
beneat h t he plasma membr ane.
3. Act in and myosin cont r act t o pr oduce t he const r ict ion.
4. Plant s have cell walls and t he cyt oplasm divides differ ent ly.
• Aft er t he spindle br eaks down, vesicles fr om t he Golgi appar at us appear in t he
equat or ial r egion.
• The vesicles fuse t o for m a new plasma membr ane, and t he cont ent s of t he
vesicles combine t o for m t he cell plat e, which is t he beginning of t he new cell
wall.
5. Or ganelles and ot her cyt oplasmic r esour ces do not need t o be dist r ibut ed equally in
daught er cells, as long as some of each ar e pr esent in bot h new cells t o assur e addit ional
gener at ion of or ganelles as needed.
VI. Reproduction: Sexual and Asexual
1. Mit osis by r epeat ed cell cycles can give r ise t o vast number s of ident ical cells.
2. Meiosis r esult s in just four pr ogeny, which usually do not fur t her duplicat e. The cells
can be genet ically differ ent .
Reproduction by mitosis results in genetic constancy
3. Asexual reproduct ion involves t he gener at ion of a new individual t hat is essent ially
genet ically ident ical t o t he par ent . It involves a cell or cells t hat wer e gener at ed by
mit osis.
• Var iat ion of cells is likely due t o mut at ions or envir onment al effect s.
4. S exual reproduct ion involves meiosis.
• Two par ent s each cont r ibut e one cell t hat is genet ically differ ent fr om t he par ent s.
• These cells oft en combine t o cr eat e var iet y among t he offspr ing beyond t hat
at t r ibut ed t o mut at ions or t he envir onment .
Reproduction by meiosis results in genetic diversity
5. Sexual r epr oduct ion fost er s genet ic diver sit y among pr ogeny. Two par ent s each
cont r ibut e a set of chr omosomes in a sex cell or gamet e. Gamet es fuse t o pr oduce a
single cell, t he zygot e, or fer t ilized egg. Fusion of gamet es is called fer t ilizat ion.
Cell Biology 19
6. In each r ecognizable pair of chr omosomes, one comes fr om each of t he t wo par ent s.
The member s of t he pair ar e called homologous chr omosomes and ar e similar , but
not ident ical, in size and appear ance. (An except ion for sex chr omosomes exist s in
some species.)
7. The homologous chr omosomes have cor r esponding but gener ally not ident ical genet ic
infor mat ion.
8. Haploid cells cont ain just one homolog of each pair . The number of chr omosomes in
a single set is denot ed by n. When haploid gamet es fuse in fer t ilizat ion, t hey cr eat e
t he zygot e, which is 2n, or diploid.
9. Some or ganisms have a pr edominant life cycle in a 1n (haploid) st at e. (Algae & fungi)
10. Some or ganisms have bot h a 1n veget at ive life st age and a 2n veget at ive life st age.
(Br yophyt e/pt er idophyt es)
11. In diplont ic or ganisms, which include animals, t he or ganism is usually diploid. (Some
insect s ar e except ed.)
12. Homologous chr omosomes exchange par t s and r ecombine dur ing meiosis so t hat t he
chr omosomes passed on t o gamet es ar e mixt ur es of t hose r eceived fr om t wo par ent s.
The t wo chr omosomes of a mixed homologous pair t hen segr egat e r andomly int o
haploid gamet es. This shuffling gr eat ly incr eases t he diver sit y of t he populat ion and
oppor t unit ies for evolut ion.
The number, shapes, and sizes of the metaphase chromosomes constitute the karyotype
13. It is possible t o count and char act er ize individual chr omosomes.
14. Cells in met aphase can be killed and pr epar ed in a way t hat spr eads t he chr omosomes
ar ound a r egion on a glass slide. A phot ogr aph of t he slide can be t aken, and images
of each chr omosome can be or ganized based on size, number and shape. This spr ead
is called a karyot ype.
VII. Meiosis: A Pair of Nuclear Divisions
1. Meiosis consist s of t wo nuclear divisions t hat r educe t he number of chr omosomes t o
t he haploid number .
2. The nucleus divides t wice, but t he DNA is r eplicat ed only once.
3. The funct ions of meiosis ar e t o r educe t he chr omosome number fr om diploid t o haploid,
t o ensur e each gamet e get s a complet e set , and t o pr omot e genet ic diver sit y among
pr oduct s.
4. Meiosis I is unique for t he pair ing and synapsis of homologous chr omosomes in
pr ophase I of t he fir st nuclear division. Aft er met aphase I, homologous chr omosomes
separ at e int o differ ent cells.
5. Individual chr omosomes, each wit h t wo chr omat ids, r emain int act unt il met aphase
of meiosis II (second nuclear division) is complet ed and t he chr omat ids separ at e t o
become chr omosomes.
The first meiotic division reduces the chromosome number
6. Like mit osis, meiosis I is pr eceded by an int er phase in which DNA is r eplicat ed.
Meiosis I begins wit h a long pr ophase.
20 CSIR-NET Life Sciences
7. Dur ing pr ophase I, synapse occur s: The t wo homologs ar e joined t oget her held by a
synapt onemal complex of pr ot eins. This for ms a t et r ad, or “bundle of four ,” which
consist s of t wo homologous chr omosomes wit h t wo sist er chr omat ids.
8. At a lat er point , t he chr omosomes appear t o r epel each ot her except at t he cent r omer e
and at point s of at t achment s, called ch i asm at a, which a ppea r X-sha ped. These
chiasmat a r eflect t he exchange of genet ic mat er ial bet ween homologous chr omosomes,
a phenomenon called crossing-over.
9. This cr ossing-over incr eases genet ic var iat ion by “mixing and wat ching” t he genes on
t he homologs.
10. In t he t est is cells of human males, pr ophase I t akes about a week.
11. In t he egg cells of human females, pr ophase I begins befor e bir t h in some eggs and
can cont inue for 50 year s in ot her s depending on t heir r elease in t he mont hly ovar ian
cycle.
12. Following t elophase I, in some species, t her e is a r eappear ance of nuclear envelopes.
If t his occur s, it is called int er kinesis, a st age similar t o mit ot ic int er phase, but t her e
is no r eplicat ion of genet ic mat er ial and no cr ossing-over in subsequent st ages.
The second meiotic division separates the chromatids
13. Meiosis II is similar t o mit osis.
14. One differ ence is t hat DNA does not r eplicat e befor e meiosis II. The number of
chr omosomes is t her efor e half t hat found in diploid mit ot ic cells.
15. In meiosis II, sist er chr omat ids ar e not ident ical and t her e is no cr ossing-over .
Meiosis leads to genetic diversity
16. The pr oduct s of meiosis I ar e genet ically diver se.
17. Synapsis and cr ossing over dur ing pr ophase I mix genet ic mat er ial of t he mat er nal
wit h t hat of t he pat er nal homologous chr omosomes.
18. Which member of a homologous pair segr egat es or goes t o which daught er cell at
anaphase I is simply chance.
19. Since most species of diploid or ganisms have mor e t han t wo pair s of chr omosomes,
t he possibilit ies for var iat ion in combinat ions becomes huge.
EXTRA CONCEPTS: Cell Reproduction or Cell Division
1. Cell Division was fir st st udied by St r a s b u r ge r (1875) in plant s, W. Flemmi n g (1882)
in animal cells and P r e vos t a n d Du ma s (1824) in fr og egg.
2. Any agent t hat st imulat es cell division is called mi t ogen . Temper at ur e, cyt okinin,
auxin, gibber llin, insulin, st er oids and mit ogens.
3. The cont inuat ion of species fr om one gener at ion t o next is gover ned by t wo pr ocesses;
syngamy (union of gamet es) and division of cells (meiosis and mit osis).
4. A cell divides t o have high sur face ar ea per unit of volume and high nucleocyt oplasmic
r at io. The smaller t he size of cell, mor e t he sur face ar ea and nucleo-cyt oplasmic r at io
it has.
5. Genet ic cont inuit y is due t o duplicat ion of DNA in cell division t hat occur s in S-phase.
Cell Biology 21
6. Mi t ot i c p oi s on s ar e inhibit or s of cell divisions. Azid es and cya n id es inhibit pr ophase;
colch i ci n es checks spindle for mat ion; ch a lon e s inhibit cell division in vivo and in
vit ro bot h; r i b on u cl e a s e blocks pr ophase; h e a t s h ock s pr event cell division and
Mu s t a r d gas agglut inat e all chr omosomes.
7. Animal cyt okinesis is cent r ipet al and plant cyt okinesis is cent r ifugal.
8. In fungi, spindle is for med inside nucleus (intranuclear division); nuclear membr ane
r emains int act ; nucleus divides by fur r ow (karyochoriosis ).
9. En d omi t os i s is duplicat ion of chr omosomes wit hout division of nucleus.
10. Non-disjunct ion is failur e of migr at ion of chr omat ids at anaphase; discover ed by
Br i d ges (1961).
11. Br a c h yme i os i s : It is believed by some mycologist s t ha t in some a scomycet es,
fer t ilizat ion t akes place in single celled st age r esult ing in a diploid nucleus which
t hen under goes fr ee nuclear divisions followed by pair ing (dikar yon for mat ion). These
dikar yons t hen fuse and t hus become a t et r aploid nucleus. This is ascus mot her cell.
If it has t o for m haploid ascospor es it must now under go t wo r educt ional & one
equat ional division. This is knows as b r a ch yme i os i s .
12. Ace t oca r mi n e is made by dissolving car mine dye (obt ained fr om cochineal Coccus
insect ) in acet ic acid. It gives pur ple r ed colour t o chr omosomes.
13. C-mi t os i s is colchicines induced mit osis. Col ch i ci n e is an alkaloid, obt ained fr om
under gr ound cor ms of aut umn cr ocus – Colchicum autumnale. It was discover ed by
Du s t i n (1934) and used by Bla k eslee (1937) t o induce polyploidy. Gr a n os a n is similar
t o colchicines in act ion. Bot h inhibit spindle for mat ion.
14. In Cyperus, one meiosis pr oduces one pollen gr ain inst ead of four .
15. Cell doubles in size and t hen st ops gr owing in G
1
pha se. G
1
is longest , most var iable
phase in which maximum gr owt h occur s. Cir cumst ances which induce a cell t o divide
ar ise in G
1
under t he influence of some cyt oplasmic clock. De ci s i on for ce ll d i vi s i on
a ls o occu r s h e r e .
16. Size of nucleus incr eases in int er phase; size of nucleolus incr eases in fir st four
subst ages of Pr ophase-I of meiosis.
17. Arist olochia (duck weed) has all t ypes of t et r ads.
18. Pr ot ein (hist one) for DNA synt hesis is for med in S-phase while t ubulin pr ot ein r equir ed
for spindle is synt hesized in G
2
.
19. All or ganelles (or ganoids) including cent r ioles ar e doubled in G
2
.
20. Amount of DNA doubles in S-phase.
21. Int er phase is most act ive phase followed by pr ophase. This int er phase t akes 70-95%
of t ot al t ime of cell cycle. M-phase t akes ver y less t ime.
22. Ami t os i s is called dir ect or incipient cell division and is found in yeast s, pr ot ozoans,
moner ans (pr okar yot es), car t ilage and degener at ed/old t issues.
23. Mit osis for ms 2 daught er cells t hat ar e mor phologically and genet ically similar . It
dist r ibut es chr omosomes equally bot h quant it at ively and qualit at ively. Te r m mi t os i s
was given by W. F l e mmi n g (1870). Mer ist ems, cells of bone mar r ow, base of nails
and skin ar e used t o st udy mit osis.
22 CSIR-NET Life Sciences
24. Spindle is ast r al (amphiast r al) and ar ises fr om cent r iole in animal cell and in plant
cell; it is anast r al and ar ise fr om cyt oplasmic pr ot eins by gelat ion. It consist s of
micr ot ubules made up of sulphur r ich t ubulin pr ot ein (95-97%), RNA (3-5%) and
ATPase. Sp i n d l e i s s e e n wi t h p ol a r i zi n g mi cr os cop e on l y.
25. Pr ophase is of longest dur at ion.
26. Met aphasic chr omosomes ar e least coiled. Th e s t r u ct u r e of ch r omos ome s i s b e s t
s t u d i e d a t me t a p h a s e wh i l e s h a p e o f c h r o mo s o me s i s b e s t s t u d i e d a t
a n a p h a s e.
27. An a p h a s e i s of s h or t e s t d u r a t i on ; cent r omer e divides and disjunct ion occur s her e.
About 30 molecules of ATP ar e needed t o move one chr omosome fr om equat or t o
pole.
28. Telophase is r ever se of pr ophase. Nuclear membr ane r eappear s fr om ER and r emnant s
of or iginal nuclear membr ane.
29. Me i os i s was fir st discover ed by Bover i (1892), st udied by St r a s b u r ge r (1883) and
Wi n i wa r t e r (1990); t e r m b y F a r me r a n d Moor e (1905). It is double division in
which nucleus divides t wice but chr omosomes only once. It is ant it hesis of fer t ilizat ion
a nd ha vles t he number of chr omosomes. I t ma int a ins number of chr omosomes
const ant t hr ough successive gener at ions.
30. Meiosis occur s in diploid r epr oduct ive cells (meiocyt es) at t he t ime of r epr oduct ion.
31. Ant her s of unopened young flower s and t est es of gr asshopper ar e widely used t o
st udy meiosis.
32. Zygot e n e is zipping or synapsis or pair ing of homologous chr omosomes.
33. Tet r ad for mat ion occur s at pachyt ene st age.
34. Di p l ot e n e i s of l on ge s t d u r a t i on and involves chiasmat a for mat ion ver y dist inct ly.
Cr ossing over begins at pachyt ene but chiasmat a becomes dist inct at diplot ene, hence
we can say cr ossing over occur s at diplot ene.
35. Tr a n s p or t a t i on is exchange and r ejoining of chr omat ids par t s dur ing cr ossing over .
36. In met aphase-I, migr at ing chr omosomes ar e d ya d , i.e., ea ch chr omosome ha s 2
chr omat ids.
37. Reduct ion in number of chr omosomes occur in anaphase-I but haploidy (r educt ion) in
t er ms of DNA occur dur ing anaphase-II.
38. In Trillium, anaphase-I dir ect ly ent er s int o met aphase-II.
39. Tet r ad is a gr oup of 4 haploid cells for med dur ing meiosis. It can be t et r ahedr al,
isobilat er al, linear , decussat e or T-shaped but t e t r a h e d r a l t e t r a d i s mos t common
i n p l a n t s .
40. Ki n e t och or e is a pr ot einaceous r egion of t he cent r omer e in chr omosome t o which
spindle fibr es at t ach.
41. Nucleopr ot ein complex pr esent bet ween synapsed chr omosomes is called synapt inemal
complex.
42. Kar yokinesis is division of nucleus. It was fir st st udied by Sch lei d en .
43. Chr omat ids move t owar ds t he pole at a speed of 1 µm per minut e.
44. Chiasmat a ar e t he r esult of cr ossing over and fir st obser ved by J a n s e e n s (1909).
Cell Biology 23
45. He La cells ar e human cancer cells of a pat ient Hen r i et t a La ck ; maint ained in
t issue cult ur e since 1953. They divide and double t heir number in ever y 24 hr s &
widely used in r esear ch.
46. Dur ing G2, a cell cont ains double t he amount of DNA (4n) as compar ed t o or iginal
diploid cell (2n).
47. Re p a i r of d a ma ge d DNA a l s o t a k e s p l a ce i n t h e i n t e r p h a s e.
48. In plant s, mit osis occur in mer ist emat ic t issues (shoot & r oot t ips). Root t ip is t he
most pr efer r ed r egions t o obser ve mit osis.
49. An a p h a s e i s a r a p i d p h a s e l a s t i n g on l y 2-3 mi n u t e s . It st ar t s abr upt ly. The
cent r omer e split s int o t wo; each chr omat id is pulled slowly t owar ds a spindle pole
(each chr omat id wit h own cent r omer e now becomes a separ at e single st r anded (1
DNA) chr omosome. The chr omat ids ar e moved (t owar ds t he pole t hey face) at a
speed of 1µm/minut e. The separ at ion of t he chr omat ids st ar t s at t he cent r omer es
while t he ar ms t r ail behind it . As a r esult , t he chr omosomes ar e pulled int o V, J and
T shapes.
50. The t elophase last s for an hour or so.
51. St i mu l a t i on of mi t os i s : Kinet in (6-fur t ur yl amino pur ine) incr eases t he mit ot ic
r at e in mer ist ems of Allium. At low concent r at ion, it r educes t he dur at ion of int er phase
and incr eases t he mit ot ic r at e.
52. In human males, meiosis st ar t s aft er puber t y.
53. In human females, meiosis st ar t s at t he end of 3
rd
mont h of pr enat al life. In t he fift h
mont h of pr enat al life, t he oocyt es r each t he diplot ene st age and r emain ar r est ed at
t his st age for many (About 12) year s, when ovulat ion occur s.
54. Number of meiosis required to form n number of seeds/grains = n + n/4

(for all cases
except cyper us); in cyper us, it is = n + n.
CELL DIFFERENTIATION AND CELL-CELL INTERACTION:
Specializations of Plasmalemma
They ar e of t hr ee t ypes:
(i) Out pushings (Evaginat ions) - micr ovilli, flagellar or ciliar y sheat hs, st er eocilia.
(ii) Inpushings (Invaginat ions) - por es, mesosomes, lomasomes and t r ansfer cells.
(iii) J unct ional Complexes. They ar e connect ions bet ween adjacent cells, acr oss int er vening
space of 15 - 20 nm widt h which is oft en filled wit h t issue fluid. Cement ing mat er ial is
called adher enes, fusion as occludens, spot as macula and st r ip as zonula. Common
junct ional complexes ar e plasmodesmat a, gap junct ions, int er digit at ions, int er cellular
br idges, t ight junct ions, desmosomes and t er minal bar s.
1. Mi c r ovi l l i (Singular -Micr ovillus). They ar e numer ous (upt o 3000) fine plasmalemma
evaginat ions (each O’ 6 - 0,8 ,um long and 0.1 ,um in diamet er ) which gives st r iat ed or
br ush bor der appear ance under opt ical micr oscope. Micr ovilli ar e suppor t ed int er nally
by micr o filament s. Ext er nally t hey possess glycocalyx. Ar eas in bet ween t he micr ovilli
ar e specialised for absor pt ion. Sur face ar ea is incr eased sever al t imes, e.g., int est inal
epit helium, hepat ic cells, convolut ed r egions of r enal t ubules, lining of gall bladder
and ut er us.
24 CSIR-NET Life Sciences
2. St e r e oci l i a . Nonmot ile elongat ed evaginat ions of plasma membr ane, secr et or y or
sensor y, e.g., macula, cr ist a, epididymus. Tr ue cilia and flagella ar e cover ed by plasma
membr ane sheat hs. Evaginat ions also occur dur ing for mat ion of phagocyt ic vesicles.
3. P or e s . At places plasma membr ane is connect ed wit h endoplasmic r et iculum for ming
por es leading t o channels of E.R. Infolds also develop dur ing for mat ion of endocyt ot ic
vesicles.
4. Me s os ome . Complex infolding of plasma membr ane in bact er ia t hat is connect ed
wit h nucleoid and is believed t o help in nucleoid r eplicat ion, sept um for mat ion and
even r espir at ion. A similar infolding found in fungi is called lomasome.
5. Tr a n s fe r ce lls . Bot h cell wall and plasmalemma show infoldings. Cells ar e specialized
for solut e t r ansfer .
6. P l a s mod e s ma t a . They ar e cyt oplasmic br idges bet ween adjacent plant cells t hat
occur in ver y fine por es or pit s in t he cell wall.
7. Ga p J u n ct i on s (Nexus, Maculae Occludent es). Fine hydr ophilic channels for med by
special pr ot ein cylinder s or connexons of t wo adjacent cells. Ca
2+
is r equir ed for t heir
opening, t hey a r e ver y common.
8. I n t e r ce l l u l a r Br i d ge s . Plasma pr oject ions fr om adjacent cells t hat come in cont act
in t he int er cellular space for quick t r ansfer of st imuli.
9. I n t e r d i gi t a t i on s . Membr ane out gr owt hs of adjacent cells which fit int o one anot her ,
incr ease adher ence and sur face ar ea for exchange of mat er ials.
10. De s mos ome s (Spot Desmosomes, Maculae Adher ent es). They ar e just like welded
ar eas bet ween adjacent cells having int er cellular t hickening mat er ials, t r ansmembr ane
linker s, disc-sha ped int r a cellula r t hickening a dja cent t o ea ch membr a ne, wit h
t on ofi br i l s . De s mos ome s occu r i n e p i t h e l i a s u bj e ct e d t o d i s r u p t i on . I n
hemidesmosome, disc-shaped int r acellular t hickening occur s in one cell. Collagen
fir bils ar e found in place of int er cellular t hickening. Sept at e desmosomes possess
t r ansver se sept a in bet ween cells inst ead of int er cellular cement . Tonofibr ils ar e
absent . Sept at e desmosomes occur in inver t ebr at es.
11. Ti gh t J u n ct i on s (Zonulae Occludent es). Plasmalemmae of t wo cells fused t o for m
imper meable or occluding junct ions, e.g., epit helial cells or capillar ies and br ain cells.
Funct ion of t ight junct ions is differ ent in differ ent t issues.
12. Te r mi n a l Ba r s (= Belt Desmosomes, Zonulae Adher ent es). Desmosomes which lack
t onofibr ils and wher e discoid t hickenings ar e r eplaced by bands of micr ofilament s
and int er mediat e filament s.
CANCER AND MALIGNANT GROWTH
1. Cancer is a disease of t he body’s cells. It occur s when cells in t he body become abnor mal
and gr ow out of cont r ol. A change which makes t he gene fault y is called a mut at ion.
Some special genes, called cont r ol genes, inst r uct t he cell t o copy it s genes cor r ect ly,
and t o divide in an or der ly manner . They st op cont r olling cell division, which is
cancer .
2. Be n i gn Tu mor s : Tumor s ar ise wit h gr eat fr equency, especially in older animals and
humans, but most pose lit t le r isk t o t heir host because t hey ar e localized and of small
size. The sur face int er act ion molecules t hat hold t issues t oget her keep benign t umor
Cell Biology 25
cells, like nor mal cells, localized t o appr opr iat e t issues. A fibr ous capsule usually
delineat es t he ext ent of a benign t umor .
3. Ma l i gn a n t t u mor : In cont r ast , t he cells composing a malignant t umor , or cancer ,
expr ess some pr ot eins char act er ist ic of t he cell t ype fr om which it ar ose, and a high
fr act ion of t he cells gr ow and divide mor e r apidly t han nor mal.
4. Some malignant t umor s r emain localized and encapsulat ed, at least for a t ime; an
example is car cinoma in sit u in t he ovar y or br east .
5. Most , however , do not r emain in t heir or iginal sit e; inst ead, t hey invade sur r ounding
t issues, get int o t he body’s cir culat or y syst em, and set up ar eas of pr olifer at ion away
fr om t he sit e of t heir or iginal appear ance.
6. The spr ead of t umor cells and est ablishment of secondar y ar eas of gr owt h is called
me t a s t a s i s ; most malignant cells event ually acquir e t he abilit y t o met ast asize.
7. Thus t he major char act er ist ics t hat differ ent iat e met ast at ic (or malignant ) t umor s
fr om benign ones ar e t heir invasiveness and spr ead.
Characteristics of Cancer Cells
1. They ar e usually less well differ ent iat ed t han nor mal cells or benign t umor cells. The
pr esence of invading cells is t he most diagnost ic indicat ion of a malignancy.
2. Cancer cells can mult iply in t he absence of gr owt h-pr omot ing fact or s r equir ed for
pr olifer at ion of nor mal cells and ar e r esist ant t o signals t hat nor mally pr ogr am cell
deat h (apopt osis).
3. Cancer cells also invade sur r ounding t issues, oft en br eaking t hr ough t he basal laminas
t hat define t he boundar ies of t issues and spr eading t hr ough t he body t o est ablish
secondary areas of grow th, a process called metastasis
4. Bot h pr imar y and secondar y t umor s r equir e angiogenesis, t he r ecr uit ment of new
blood vessels, in or der t o gr ow t o a lar ge mass.
5. Cancer cells, which ar e closer in t heir pr oper t ies t o st em cells t han t o mor e mat ur e
differ ent iat ed cell t ypes, usually ar ise fr om st em cells and ot her pr olifer at ing cells
Types of Cancer
1. Ca r ci n oma : It includes t umor s of br ain, br east , skin, cer vical r egion. These ar e
der ived fr om epit helial t issue, or iginat ing fr om eit her ect oder m or endoder m. These
occur s as solid t umor s, locat ed in t he ner vous t issue on t he body sur face or associat ed
glands.
2. Sa r coma : They ar e t he cancer s of connect ive t issues, car t ilage, bone or muscles
which ar e mesoder mal in or igin.
3. Th e le u k e mi a s : A class of sar comas, gr ow as individual cells in t he blood, wher eas
most ot her t umor s ar e solid masses. (The name leukemia is der ived fr om t he Lat in
for “whit e blood”: t he massive pr olifer at ion of leukemic cells can cause a pat ient ’s
blood t o appear milky)
4. Ly mp h o ma: Lymph nodes, bone ma r r ow, liver a nd spleen pr oduces excessive
lymphocyt es. Cancer in t hem ar e called as lymphomas eg. Hodgkin’s disease.
26 CSIR-NET Life Sciences
Proto-Oncogenes and Tumor-Suppressor Genes
1. Two br oad classes of genes—pr ot o-oncogenes (e.g., ras) and t umor -suppr essor genes
(e.g., APC) play a key r ole in cancer induct ion. These genes encode many kinds of
pr ot eins t hat help cont r ol cell gr owt h and pr olifer at ion; mut at ions in t hese genes can
cont r ibut e t o t he development of cancer .
2. Most cancer s have inact ivat ing mut at ions in one or mor e pr ot eins t hat nor mally
funct ion t o r est r ict pr ogr ession t hr ough t he G
1
st age of t he cell cycle (e.g., Rb and
p16). Vir t ually all human t umor s have inact ivat ing mut at ions in pr ot eins such as p 53
t hat nor mally funct ion at cr ucial cell-cycle checkpoint s, st opping t he cycle if a pr evious
st ep has occur r ed incor r ect ly or if DNA has been damaged. Likewise, a const it ut ively
act ive Ras is found in sever al human t umor s of differ ent or igin. Thus nor mal gr owt h
cont r ol and malignancy ar e t wo faces of t he same coin.
3. An oncogene is any gene t hat encodes a pr ot ein able t o t r ansfor m cells in cult ur e or
t o induce cancer in animals.
4. Of t he many known oncogenes, all but a few ar e der ived fr om nor mal cellular genes
(i.e., pr ot o-oncogenes) whose pr oduct s pa r t icipa t e in cellula r gr owt h-cont r olling
pat hways. For example, t he ras gene is a pr ot o-oncogene t hat encodes an int r acellular
signal-t r ansduct ion pr ot ein;
5. Conver sion, or act ivat ion, of a pr ot o-oncogene int o an oncogene gener ally involves a
gain-of-function mut at ion.
6. Tumor -suppr essor genes gener ally encode pr ot eins t hat in one way or anot her inhibit
cell pr olifer at ion. Loss of one or mor e of t hese “br akes” cont r ibut es t o t he development
of many cancer s.
7. Five br oad classes of pr ot eins ar e gener ally r ecognized as being encoded by t umor -
suppr essor genes:
• Int r acellular pr ot eins, such as t he p16 cyclin-kinase inhibit or , t hat r egulat e or
inhibit pr ogr ession t hr ough a specific st age of t he cell cycle
• Recept or s for secr et ed hor mones (e.g., t umor der ived gr owt h fact or ?) t hat funct ion
t o inhibit cell pr olifer at ion
• Checkpoint -cont r ol pr ot eins t hat ar r est t he cell cycle if DNA is damaged or
chr omosomes ar e abnor mal
• Pr ot eins t hat pr omot e apopt osis and Enzymes t hat par t icipat e in DNA r epair
Some of the characteristics of Oncogenes and tumour suppressor genes can be
summarized as follows
1. Dominant gain-of-funct ion mut at ions in pr ot o-oncogenes and r ecessive loss-of-funct ion
mut at ions in t umor -suppr essor genes ar e oncogenic.
2. Among t he pr ot eins encoded by pr ot o-oncogenes ar e posit ive-act ing gr owt h fact or s
and t heir r ecept or s, signal-t r ansduct ion pr ot eins, t r anscr ipt ion fact or s, and cell-cycle
cont r ol pr ot eins.
3. An act ivat ing mut at ion of one of t he t wo alleles of a pr ot o-oncogene conver t s it t o an
oncogene, which can induce t r ansfor mat ion in cult ur ed cells or cancer in animals.
4. Act ivat ion of a pr ot o-oncogene int o an oncogene can occur by point mut at ion, gene
amplificat ion, and gene t r anslocat ion.
Cell Biology 27
5. The fir st r ecognized oncogene, v-src, was ident ified in Rous sar coma vir us, a cancer -
causing r et r ovir us. Ret r ovir al oncogenes ar ose by t r ansduct ion of cellular pr ot o-
oncogenes int o t he vir al genome and subsequent mut at ion.
6. The fir st human oncogene t o be ident ified encodes a const it ut ively act ive for m of
Ras, a signal-t r ansduct ion pr ot ein. This oncogene was isolat ed fr om a human bladder
car cinoma.
7. Slow-act ing r et r ovir uses can cause cancer by int egr at ing near a pr ot o-oncogene in
such a way t hat gene t r anscr ipt ion is act ivat ed cont inuously and inappr opr iat ely.
8. Tumor -suppr essor genes encode pr ot eins cell cycle if DNA is damaged or chr omosomes
ar e abnor mal, r ecept or s for secr et ed hor mones t hat funct ion t o inhibit cell pr olifer at ion,
pr ot eins t hat pr omot e apopt osis, and DNA r epair enzymes.
9. Inher it ed mut at ions causing r et inoblast oma led t o t he ident ificat ion of RB, t he fir st
t umor -suppr essor gene t o be r ecognized.
IMMUNE RESPONSE
1. Immunit y means pr ot ect ion fr om disease and especially infect ious disease. Cells and
m olecules involved in such protection constitute the i mmu n e syst em and t he response
t o int r oduct ion of a for eign agent is known as t he i mmu n e r e s p on s e.
2. Not all immune r esponses pr ot ect fr om disease; some for eign agent s, such as t he
a l l e r ge n s found in house dust mit e, cat dander or r ye gr ass pollen, cause disease as
a consequence of inducing an immune r esponse.
3. Likewise some individuals mount immune r esponses t o t heir own t issues as if t hey
wer e for eign agent s. Thus, t he immune r esponse can cause t he a u t oi mmu n e diseases
common t o ma n such a s mult iple scler osis, dia bet es, r heuma t oid a r t hr it is or
myast henia gr avis.
4. Most individuals do not suffer fr om aut oimmune disease because t hey have developed
t ol e r a n ce t owar ds t heir own (s e lf) t issues.
Innate (or natural) immunity: This is made up of several components.
1. P h ys i ca l b a r r i e r s a r e t h e fi r s t l i n e of d e fe n s e a ga i n s t i n fe ct i on . The skin and
mucous membr anes pr ovide a cont inous sur face which must be br eached and back
t his up wit h mechanical pr ot ect ion t hr ough cilia and mucous.
2. P h ys i ol ogi ca l fa ct or s such as pH, t emper at ur e and oxygen t ension limit micr obial
gr owt h. The acid envir onment of t he st omach combined wit h micr obial compet ion
fr om t he commensal flor a inhibit s gut infect ion.
3. P r ot e i n s e cr e t i on s int o ext er nal body fluids such as lysozyme also help r esist
invasion. Soluble fact or s wit hin t he body such as comp l e me n t , i n t e r fe r on s and
collect ins and ot her “br oadly specific” molecules such as C-r eact ive pr ot ein ar e of
consider able impor t ance in pr ot ect ion against infect ion.
4. P h a gocyt i c ce l l s ar e cr it ical in t he defense against bact er ial and simple eukar yot ic
pa t h ogen s . Ma c r o p h a g e s and P o l y mo r p h o n u c l e a r l e u c o c y t e s (P MN) ca n
r ecognise bact er ial and yeast cell walls t hr ough br oadly specific r ecept or s (usually for
ca r bohydr a t e st r uct ur es) a nd t his r ecognit ion is gr ea t ly enha nced by a ct iva t ed
complement (opsonin) [as well as by specific ant ibody].
28 CSIR-NET Life Sciences
5. Acu t e I n fl a mma t i on : The acut e inflammat or y r esponse which has been descr ibed
in pr evious lect ur es is a key par t of t he innat e immune syst em. Many infect ions,
especia lly wher e sma ll wounds a r e t he r out e of ent r y, a r e elimina t ed by t he
combinat ion of complement and r ecr uit ment of phagocyt es, which flow fr om t he acut e
inflammat or y r esponse.
A defining aspect of t he innat e immune syst em is t hat it carries no memory of an
encount er wit h a foreign organism.
What is an antigen?
1. An a n t i gen is defined as “anyt hing t hat can be bound by an ant ibody”. This can be an
enor mous r ange of subst ances fr om simple chemicals, sugar s, small pept ides t o complex
pr ot ein complexes such as vir uses.
2. The small ant igens ar e not , however i mmu n oge n i c in t hemselves and need t o be
coupled t o a ca r r i er t o elicit an immune r esponse. Such small ant igens ar e r efer r ed
t o as h a p t e n s .
3. Requir ement of Ant igen t o ca use immune r esponse a r e: Non-self, Complex in
st r uct ur e, should mor e t hen 5 KD in size and must have at least on epit ope.
4. In fact ant ibodies int er act specifically wit h r elat ively small par t s of molecules. These
ar e known as ant igenic d e t e r mi n a n t s or e p i t op e s .
5. Somet imes t he epit ope is composed of a st r ing of amino acids as might be found in a
shor t pept ide, such epit opes ar e said t o be li n e a r . Ot her epit opes ar e for med by mor e
complex 3-dimensional st r uct ur es pr esent only as par t of a nat ive pr ot ein, such epit opes
ar e called con for ma t i on a l .
Adaptive immunity
1. The second level of defence incr ea ses in st r engt h a nd effect iveness wit h ea ch
encount er . The for eign agent is r ecognised in a s p e ci fi c manner and t he immune
syst em a cqu i r es memor y t owar ds it .
1°challenge 2°boost
New challenge
response to new
a
n
t
i
b
o
d
y

r
e
s
p
o
n
s
e
time in days
0 5 10 15 20 25 30 40 45 35
Specific memory is the hallmark
of the adaptive immune response
Cell Biology 29
2. The fir st encount er wit h an ant igen is known as t he p r i ma r y r e s p on s e. Re-encount er
wit h t he same ant igen causes a s e con d a r y r e s p on s e t hat is mor e r apid and power ful.
3. Acquir ed immunit y is a useful evolut ionar y adapt at ion because it impr oves t he efficacy
of t he innat e immune r esponse by focusing t he r esponse t o t he sit e of invasion/
infect ion as well as pr oviding addit ional effect or mechanisms t hat ar e unique t o
l ymp h ocyt e s .
4. The differ ence bet ween innat e and acquir ed immunit y lies in t he a n t i gen sp eci fi ci t y
of lymphocyt es. This pr oper t y is confer r ed upon lymphocyt es by t he expr ession of cell
sur face r ecept or s t hat r ecognise discr et e par t s of t he ant igen known as a n t i ge n i c
e p i t op e s .
5. The cell sur face r ecept or of B l ymp h ocyt e s , (der ived and mat ur e in Bon e ma r r ow
in mammals or t he Bu r s a of fa b r i ci u s in chickens) is an i mmu n oglob u li n molecule
which, when secr et ed by t he B cell, is known as an a n t i b od y.
6. Immunit y pr ovided by Immunoglobulins (ant ibodies) is t er med as humor al immunit y
(Humor =fluid/blood)
Antibodies work in three ways
1. Aggl u t i n a t i on : Ant ibodies bind t o ant igens t o pr oduce lar ge insoluble complexes,
which r ender t hem har mless and facilit at e t heir dest r uct ion by ot her cells of t he
immune syst em.
2. Op s on i a t i on : I
g
G molecules coat t he sur face of ant igens and t his st imulat es t heir
r ecognit ion and digest ion by phagocyt es.
3. Comp l e me n t me d i a t e d ce l l l ys i s : Complement syst em is a gr oup of enzymes.
They ar e t r igger ed by I
g
M - I
g
G bound t o t he sur face of for eign cells. The act ivit y of
some of t hese enzymes leads t o t he for mat ion of por es on t he plasma membr ane of
t he invading cell causing t hem t o lyse.
St r u ct u r e of An t i b od y: Ant ibodies have t wo ends. One end int er act s wit h t he ant igen
(t h e v a r i a b l e pa r t ) l ea vi n g t h e ot h er (c o n s t a n t ) en d fr ee t o in t er a ct wit h t h e
immunoglobulin r ecept or s on t hese cells. Dur ing an immune r esponse, a complex lat t ice
of int er linked ant igens and ant ibodies, known as an i mmu n e comp le x , will pr esent an
30 CSIR-NET Life Sciences
ar r ay of const ant r egions which can act ivat e t he var ious cells ment ioned above t hr ough
ligat ion of t heir immunoglobulin r ecept or s.
1. An t i bod i es : Immunoglycopr ot eins secr et ed by B-lymphocyt es in r esponse t o ant igens.
They ar e Y shaped molecule made up of t wo heavy and t wo light chains [Kappa or
lambda eit her one of t hem. They occur in t he r at io of 2:1 in human’s er a]. The ant igen
combining sit e of molecule is aminot er minus. It is composed of bot h L and H chains.
The r egions in t he L and H var iable por t ion t hat act ually combines wit h t he ant igenic
det er minant s ar e called hot spot s or iodit ypes. The car boxyt er minus of light chain
has a small const ant r egion. The heavy chain also has t wo t o t hr ee const ant r egions
at it s car boxy t er minus. Depending upon differ ent t ypes of const ant r egions, five
isot ypes of I
g
have been ident ified: I
g
G, I
g
A, I
g
M, I
g
D, I
g
E
I
g
G I
g
A I
g
M I
g
D I
g
E
Sediment at ion Coefficient 7 7 19 7 8
Molecular weight 150,000 160,000 900,000 180,000 190,000
(2 Chains) (5 Chains)
of I
g
A
Ser um concent r at ion (mg / ml) 12 2 1.2 0.03 0.00004
Half life (days) 23 6 10 2.8 2.3
Car bohydr at e % 45 42 80 75 50
Complement fixat ion Å – + – –
Placent al t r anspor t Å – – – –
Pr esent in br east milk + + – – –
Select ive secr et ion of
ser omucous glands – + – – –
Hea t st a bilit y + + + + –
2. Lymp h ocyt e s : Lymphocyt es t hat ar e cr it ical for immune r eact ions ar e of t wo t ypes:
B-cells and T-cells. Bot h develop fr om st er n cells locat ed in t he liver of t he foet us and
in t he bone mar r ow cells of t he adult . Those t hat migr at e t o t he t hymus, differ ent iat e
under it s influence and ar e known as “T-cells”; while t hose t hat cont inue t o r emain
in t he bone mar r ow ar e called “B-cells”. The young lymphocyt es migr at e t o t hymus
and lat er on lymphoid t issues such as spleen lymph nodes and t onsils wher e t hey
under go final mat ur at ion.
HUMORAL AND CELL MEDIATED IMMUNE RESPONSES
1. Hu mor a l i mmu n i t y pr oduces ant ibodies in r esponse t o t oxins (ex bee venom), fr ee
bact er ia, and vir uses pr esent in t he body fluids.
2. "Humor ” is medieval t er m for body fluids. Her e it r efer s t o t he fluid of t he blood and
t he lymph.
Cell Biology 31
3. Ant ibodies t o t hese t ypes of ant igens ar e synt hesized by B- lymphocyt es and t hen
secr et ed as soluble pr ot eins which cir culat e t hr ough t he body in blood plasma and
lymph.
4. Ce ll-me d i a t e d i mmu n i t y is t he r esponse t o int r acellular bact er ia and vir uses, fungi,
pr ot ozoans, wor ms, t r ansplant ed t issues, and cancer cells.
5. Depends on t he dir ect act ion of var ious t ypes of T-lymphocyt es r at her t han ant ibodies.
6. T-lymphocyt es differ ent iat e int o 4 differ ent var iet ies of T-cells.
• Helper /inducer T-cells (Links Cell meduat ed and Humor al Immunit y)
• Suppr essor T cells (Suppr esses t he immune r esponse when ant igens ar e disposed
off)
• Cyt ot oxic T cells or killer cells (Effect or cells of cell mediat ed immunit y)
• Memor y T-cells.
7. “Memor y B and T cells” ar e cells t hat have been exposed t o an ant igen and ar e r eadily
conver t ed t o “Effect or cells” by a lat er encount er wit h t he same ant igen. Unlike
ot her lymphocyt es, t hey per sist in t he body for mont hs or even year s.
8. The killer T-cells dir ect ly at t ack and dest r oy ant igens. They move t o t he sit e of
invasion and pr oduce chemicals t hat at t r act phagocyt es. Helper T-cells act t o st imulat e
ant ibody pr oduct ion by B-cells, while suppr essor T-cells suppr esses t he t ot al immune
syst em fr om at t acking t he body’s own cells.
9. Th ymu s -I n d e p e n d e n t An t i ge n s : Some bact er ial polysacchar ides can act ive B-cells.
Ant igens capable of doing so ar e called t hymus independent ant igens because t hese
ant igens do not r equir e t he cooper at ion of Helper T-cells for act ivat ing B-cells.
Thymus dependent ant igens; many ant igens r equir e t he cooper at ion of T-cells for
act ivat ing B-cells. The ant igens pr esent on or gan t r ansplant ar e t hymus dependent
ant igens. So a per son whose t hymus gland is r emoved is not able t o r eject or gan
t r ansplant .
10. MHC ar e major hist ocompat ibilt y complex (glycopr ot eins) pr ensent on cells which
pr esent pr ocessed int r acellular ant igen t o T-Lymphocyt es. They ar e also called as
Human Leucocyt e Ant igen (HLA) in humans.
11. MHC-I is pr esent on all nucleat ed cells and ar e r ecognized by CD 8
+
ar ms of T-
Cyt ot oxic cells and pr esent pr ocessed ant igen t o T-cyt ot oxic cells while MHC-I I ar e
pr esent on T & B lymphocyt es, dendr it ic cells and mast cells and r ecognized via CD
4
+
ar m pr esent on T-Helper cell.
12. Lymp h ok i n e s : They ar e a family of soluble chemical mediat or r eleased by helper T-
cells. Lymphokines have t wo br oad funct ions. One gr oup of lymphokines is concer ned
wit h gr owt h and differ ent iat ion of B and T cells. The ot her gr oup helps phagocyt osis
by at t r act ing phagocyt ic cells when r equir ed t hr ough chemot axis and by act ivat ing
t he phagocyt ic cells.
VACCINES
The int r oduct ion of vaccinat ion has been one of t he most decisive advances leading t o t he
dr amat ic downwar d t r end in t he incidence of many vir al diseases.
32 CSIR-NET Life Sciences
The pr inciple of vaccinat ion is t o induce a “p r i me d ” s t a t e in t he vaccinat ed subject so
t hat , following exposur e t o a pat hogen, a r apid s e con d a r y i mmu n e r e s p on s e is gener at ed
leading t o t he acceler at ed eliminat ion of t he or ganism and pr ot ect ion fr om clinical disease.
Success depends on t he gener at ion of me mor y T a n d B ce l l s and t he pr esence in t he ser um of
n e u t r a l i zi n g a n t i b od y.
At t r i b u t e s of a good va cci n e
• Ab i l i t y t o e l i ci t t h e a p p r op r i a t e i mmu n e r e s p on s e for t he par t icular pat hogen:
Tuber culosis—cell mediat ed r esponse
Most bact er ial and vir al infect ions—ant ibody
• Lon g t e r m p r ot e ct i on ideally life-long
• Sa fe t y vaccine it self should not cause disease
• St a b l e r et a i n i mmu n ogen i ci t y, des pi t e a dver s e s t or a ge con di t i on s pr i or t o
administ r at ion
• I n e xp e n s i ve
Typ es of Va cci n e: Vaccines in gener al use includes: LIVE vaccines; and KILLED vaccines
A. Live Vaccines
1. Live attenuated organisms
Or ganisms whose vi r u le n ce h a s b e e n a r t i fi ci a lly r e d u ce d by in vit ro cult ur e
under adver se condit ions, such as r educed t emper at ur e. This r esult s in t he select ion
of mu t a n t s which r eplicat e poor ly in t he human host and ar e t her efor e of r educed
vir ulence. Replicat ion of t he vaccine st r ain in t he host r epr oduces many of t he feat ur es
of wild t ype infect ion, wi t h ou t ca u s i n g cl i n i ca l d i s e a s e . Most successful vir al
vaccines belong t o t his gr oup.
The immune r esponse is usually good—when t he vir us r eplicat es in t he host cells,
bot h a n t i b od y as well as ce l l me d i a t e d i mmu n e r e s p on s e s ar e gener at ed and
immunit y is gener ally long lived. Oft en, only a s i n gl e d os e is needed t o induce long
t er m immunit y.
P ot e n t i a l d r a wb a ck s t o t hese vaccines include: t he danger of r ever sion t o vir ulence
and t he possibilit y of causing ext ensive disease in immuno-compr omised individuals.
2. Heterologous vaccines
Closely r elat ed or ganism of lesser vir ulence, which shar es many ant igens wit h t he
vir ulent or ganism. The vaccine st r ain r eplicat es in t he host and induces an immune
r esponse t hat cr oss r eact s wit h ant igens of t he vir ulent or ganism. The most famous
example of t his t ype of vaccine is vaccinia vir us: Bot h cowpox vir us and vaccinia
vir us a r e closely r ela t ed t o va r iola vir us, t he ca usit ive a gent of sma llpox. The
eight eent h cent uar y physician, Edwar d J enner obser ved t hat milkmaids who had
been infect ed wit h cowpox vir us wer e immune t o smallpox. Widespr ead use of vaccinia
vir us as a vaccine has lead t o t he wor ld-wide er adicat ion of smallpox.
3. Live recombinant vaccines
It is possible, using genet ic engineer ing, t o int r oduce a gene coding for an immunogenic
pr ot ein fr om one or ganism int o t he genome of anot her (such as vaccinia vir us). The
or ganism expr essing a for eign gene is called a r ecombinant . Following inject ion int o
Cell Biology 33
t he subject , t he r ecombinant or ganism will r eplicat e and expr ess sufficient amount s
of t he for eign pr ot ein t o induce a specific immune r esponse t o t he pr ot ein.
B. Killed (inactivated) vaccines
1. When safe live vaccines ar e not available, eit her because at t enuat ed st r ains have not
been developed or else because r ever sion t o wild t ype occur s t oo r eadily, it may be
possible to use an i n a ct i va t e d pr epar at ion of t he vir ulent or ganism t o immunize t he
host .
2. The or ganism is pr opagat ed in bulk, in vit ro, and inact ivat ed wit h eit her bet a-
pr opiolact one or for maldehyde. These vaccines ar e not infect ious and ar e t her efor e
r elat ively safe. However , t hey ar e usually of lower immunogenicit y and mult iple
doses may be needed t o induce immunit y. In addit ion, t hey ar e usually expensive t o
pr epar e.
Subcellular fr act ions
3. When pr ot ect ive immunit y is known t o be dir ect ed against only one or t wo pr ot eins
of an or ganism, it may be possible t o use a pur ified pr epar at ion of t hese pr ot eins as a
vaccine. The or ganism is gr own in bulk and inact ivat ed, and t hen t he pr ot ein of
int er est is pur ified and concent r at ed fr om t he cult ur e suspension. These vaccines ar e
sa fe a nd fewer loca l r ea ct ions occur a t t he inject ion sit e. However , t he sa me
disadvant ages of poor immunogenicit y and t he need for mult iple boost er s apply.
C. Recombinant proteins
Immunogenic pr ot eins of vir ulent or ganisms may be synt hesized ar t ificially by int r oducing
t he gene coding for t he pr ot ein int o an expr ession vect or , such as E-coli or yeast s. The pr ot ein
of int er est can be ext r act ed fr om lysat es of t he expr ession vect or , t hen concent r at ed and pur ified
for use as a vaccine. The only example of such a vaccine, in cur r ent use, is t he hepat it is B
vaccine.
D. DNA Vaccines
DNA vaccines ar e at pr esent exper iment al, but hold pr omise for fut ur e t her apy since t hey
will evoke bot h humor al and cell-mediat ed immunit y, wit hout t he danger s associat ed wit h live
vir us vaccines.
The gene for an ant igenic det er minant of a pat hogenic or ganism is inser t ed int o a plasmid.
This genet ically engineer ed plasmid compr ises t he DNA vaccine which is t hen inject ed int o
t he host . Wit hin t he host cells, t he for eign gene can be expr essed (t r anscr ibed and t r anslat ed)
fr om t he plasmid DNA, and if sufficient amount s of t he for eign pr ot ein ar e pr oduced, t hey will
elicit an immune r esponse.
E. Vaccines in general use
1. Mea sles : Live at t enuat ed vir us gr own in chick embr yo fibr oblast s, fir st int r oduced
in t he 1960’s. In developed count r ies, t he vaccine is administ er ed t o all childr en in
t he second year of life (at about 15 mont hs). If t he vaccine is administ er ed t oo ear ly,
however , t her e is a poor t ake r at e due t o t he int er fer ence by mat er nal ant ibody. For
34 CSIR-NET Life Sciences
t his r eason, when vaccine is administ er ed befor e t he age of one year , a boost er dose
is r ecommended at 15 mont hs.
2. Mu mp s : Live at t enuat ed vir us developed in t he 1960’s.In fir st wor ld count r ies it is
administ er ed t oget her wit h measles and r ubella at 15 mont hs in t he MMR vaccine.
3. Ru b e l l a : Live at t enuat ed vir us. Rubella causes a mild febr ile illness in childr en, but
if infect ion occur s dur ing pr egna ncy, t he foet us ma y develop sever e congenit a l
abnor malit ies.
4. P ol i o: Two highly effect ive vaccines cont aining all 3 st r ains of poliovir us ar e in gener al
use:
Thek i l l e d vi r u s va cci n e (Salk, 1954) is used mainly in Sweden, Finland, Holland
and Iceland.
The li ve a t t e n u a t e d or a l p oli o va cci n e (Sabin, 1957) has been adopt ed in most
par t s of t he wor ld; it s chief advant ages being: low cost , t he fact t hat it induces mucosal
immunit y and t he possibilit y t hat , in poor ly immunized communit ies, vaccine st r ains
might r eplace cir culat ing wild st r ains and impr ove her d immunit y.
Th e i n a ct i va t e d Sa l k va cci n e i s r e comme n d e d for ch i l d r e n wh o a r e
i mmu n os u p p r e s s e d .
5. He p a t i t i s B: Two va ccines a r e in cur r ent use: a ser um der ived va ccine a nd a
r ecombinant vaccine. Bot h cont ain pur ified pr epar at ions of t he hepat it is B sur face
pr ot ein. The ser um der ived vaccine is pr epar ed fr om hepat it is B sur face pr ot ein,
pur ified fr om t he ser um of hepat it is B car r ier s. This pr ot ein is synt hesised in vast
excess by infect ed hepat ocyt es and secr et ed int o t he blood of infect ed individuals. A
second vaccine, pr oduced by r ecombinant DNA t echnology, has since become available.
Thr ee doses ar e given; at 6, 10, and 14 weeks of age. As wit h any killed vir al vaccines,
a boost er will be r equir ed at some int er val (not yet det er mined, but about 5 year s) t o
pr ovide pr ot ect ion in lat er life fr om hepat it is B infect ion as a vener eal disease.
5. He p a t i t i s A: A vaccine for hepat it is A has been developed fr om for malin-inact ivat ed,
cell cult ur e-der ived vir us.
6. Va r i ce l l a -Zos t e r vi r u s : A live at t enuat ed st r ain of var icella zost er vir us has been
developed.
DOSAGE COMPENSATION
Dosage Compensation by X-inactivation in female mammals
1. How does an or ganism compensat e for t he fact t hat some individuals have a double
dosage of sex-linked genes while ot her s have only one? In female mammals, most
diploid cells have only one fully funct ional X chr omosome.
2. The explanat ion for t his pr ocess is known as t he Lyon hypot hesis, pr oposed by t he
Br it ish genet icist Mar y F. Lyon. In females, each of t he embr yonic cells inact ivat es
one of t he t wo X chr omosomes. The inact ive X chr omosome cont r act s int o a dense
object called a Bar r body.
Cell Biology 35
3. Ba r r b od y = Locat ed inside t he nuclear envelope, it is a densely st aining object t hat
is an inact ivat ed X chr omosome in female mammalian cells. Most Bar r body genes
ar e not expr essed. They ar e r eact ivat ed in gonadal cells t hat under go meiosis t o for m
gamet es. Female mammals ar e a mosaic of t wo t ypes of cells, t hose wit h an act ive
mat er nal X and t hose wit h an act ive pat er nal X. Which of t he t wo Xs will be inact ivat ed
is det er mined r andomly in embr yonic cells. Aft er an X is inact ivat ed, all mit ot ic
descenda nt s will ha ve t he sa me ina ct ive X. As a consequence, if a fema le is
het er ozygous for a sex-linked t r ait , about half of her cells will expr ess one allele and
t he ot her cells well expr ess t he alt er nat e allele. Examples of t his t ype of mosaicism
ar e color at ion in calico cat s and nor mal sweat gland development in humans. A woman
who is het er ozygous for t his t r ait has pat ches of nor mal skin and pat ches of skin
lacking sweat glands.
4. X chr omosome inact ivat ion is associat ed wit h DNA me t h yl a t i on . Me t h yl gr ou p s
(-CH
3
) a t t a ch t o cyt os i n e, one of DNA’s nit r ogenous bases. Bar r bodies ar e highly
met hylat ed compar ed t o act ively t r anscr ibed DNA. What det er mines which of t he
t wo X chr omosomes will be met hylat ed? A r ecent ly discover ed gene, XI ST is act ive
only on t he Bar r body. The pr oduct of t he XIST gene, X-inact ive specific t r anscr ipt , is
an RNA; mult iple copies of XIST at t ach t o t he X chr omosome inact ivat ing it .
Dosage Compensation by X-hyper activation in Male of Drossophila
1. In Dr ossophila Dosa ge compensa t ion is a chieved by hyper a ct iva t ion of single x
chr omosome pr esent in males. The hyper act ivit y is polyt enizat ion of X chr omosome
and hypoacet ylat ion of Hist ones at t ached t o DNA. So Genes ar e always accessible for
t r anscr ipt ion.
SEX DETERMINATION
1. Chr omosomal Basis of Sex det er minat ion:
• XX-XY is t hat t ype of sex det er minat ion in which Y det er mines sex of maleness.
It is found in mammals, sever al insect s.
• XX-XO t ype in which O det er mines sex of ma leness e.g., bugs, cockr oa ch,
gr asshopper , r oundwor ms.
• ZZ-ZW t ype – W det er min es s ex of fema len es s e.g., bir ds, r ept iles, fish,
silkwor m.
• ZO-ZZ t ype – O det er mines sex of femaleness. It is opposit e t o XX-XO t ype e.g.,
but t er flies, mot h, pigeon, ducks.
2. Ha p l oi d y- d i p l oi d y e.g., ant s, wasps, bees. Males ar e haploid and females ar e diploid.
3. Gyn a n d r omor p h is a sex mosaic (an individual wit h one half of t he body male and
t he ot her half female). These ar e common in silk mot h and Drosophila. A gynander
may be male or female wit h pat ches of t issues of ot her sex on it . Gynandr omor phism
is developed due t o n on -d i s j u n ct i on of X-ch r omos ome /a n e u p l oi d y.
36 CSIR-NET Life Sciences
4. Gen i c Ba la n ce Th eor y for sex det er minat ion in Dr ossophilla was given by Br i d ges
(on t he basis of r at io of number of X chr omosome t o set s of aut osomes)
Ch r omosome
Con st it u t ion X/A r a t i o Sex i n d ex
AA + XXX 3/2 = 1.50 Super
AA + XX 2/2 = 1.00 Nor mal
AAA + XXY 2/3 = 0.67 Int er sex
AA + XY 1/2 = 0.50 Nor mal
AA + X 1/2 = 0.50 Nor mal
AAA + XY 1/3 = 0.33 Super
5. It was concluded t hat X/A r at io of > 1.0 expr esses super femaleness, 1.0 femaleness,
1.0 femaleness, below 1.0 and above .05 int er sexes, maleness and <0.5 super maleness.
6. Y-chr omosome is male det er miner in man but not in Drossophila.
7. Envir onment al cont r ol of sex det er minat ion occur s in Tur t les, Cr ocodiles, Cer t ain
lizar ds and Alligat or s.
8. Hor monal Sex det er minat ion occur s in Scr ew’s Cock/ Hen, Fr ee Mar t in in cat t le,
Bonnelia
Practice Test Paper-I
1. In a cell if acidit y of Lysome is lost , t hen loss of:
(a) Phagocyt osis of invading bact er ia
(b) Elevat ed phosphat ase level
(c) Glycogen degr adat ion
(d) No major effect
2. Bact er ial genomes is pr event ed by it s own endonucleases by-
(a) Met hylat ion at r est r ict ion sit es
(b) Immune mechanism
(c) Nuclease r esist ant genome
(d) Ar e not much effect ive on bact er ial genome
3. The funct ion of macr ophages is t o-
(a) Enzyme Secr et ion
(b) Engulf Cell or ganelles
(c) Engulf For eign Mat er ial
(d) Kills Invading Bact er ia
4. The differ ence which dist inguish pr okar yot ic cell fr om eukar yot ic is-
(a) ER (b) Mesosome
(c) Nuclear Membr ane (d) Plasma membr ane
5. Ext r a nuclear genet ic mat er ial is found-
(a) Ribosome (b) ER
(c) Chlor oplast (d) Cent r iole
6. The acr osome of t he sper m is for med fr om t he
(a) mit ochondr ia (b) cent r osome
(c) lysomome (d) golgi bodies
7. Holiday junct ion is obser ved dur ing:
(a) Mit osis (b) Int er phase
(c) Recombinat ion (d) DNA Repair
8. A car et enoid less mut ant plant was gr own under nor mal sunlight t hen-
(a) Incr eased phot osynt hesis r at e
(b) Incr eased chlor ophyll synt hesis
(c) Reduced phot or espir at ion
(d) Incr eased chlor ophyll oxidat ion and necr osis
38 CSIR-NET Life Sciences
9. A C3 must ar d plant was gr own at 300 ppm of CO
2
in 14 h light and 10 h dar k cycles,
it was t r ansfer r ed t o 1000 ppm CO
2
. This will lead t o (ot her envir onment al par amet er s
r emaining ident ical)-
(a) Incr eased phot osynt hesis (b) Decr eased Phot osynt hesis
(c) Incr ease in Respir at ion (d) No Change
10. Pr esence of AIDS vir us cannot be det ect ed by-
(a) ELISA (b) West er n blot t ing
(c) Nor t her n Blot (d) Assay of full-lengt h ds DNA
11. Which par t of t r anslat ional modificat ion of pr ot eins does not occur in lumen of ER-
(a) Glycosylat ion
(b) Ubiquit nat ion
(c) Confor mat ion folding & for mat ion of quat er nar y st r uct ur e
(d) For mat ion of Disulphide bonds
12. Fr eshly br oken chr omosome ends ar e st icky & t end t o fuse, however ends of int act
chr omosomes ar e st able. Their st abilit y is due t o pr esence of-
(a) Cent r omer es
(b) Telomer es
(c) Special membr ane ar ound chr omosomes
(d) Kinet ochor es
13. Plant cell wall is gener ally made up of-
(a) Cellulose and pect in (b) Cellulose
(c) Chit in (d) Mur in
14. Which one of t he following is cor r ect for st r uct ur e of cell wall of fungi and Bact er ia?
(a) Bot h have glycopept ide
(b) Bot h ar e made up of N-acet ylglucasamine
(c) Bot h ar e made up of mur in
(d) Bot h ar e made up of chit in
15. Nucleus is absent in-
(a) Sieve t ube (b) cambium
(c) Phloem par enchyma (d) None of t hese
16. Among t he following which is t r ue cell accor ding cell t heor y-
(a) Vir us (b) Moner ans
(c) Pr ot est ans (d) Bact er ia
17. The char act er ist ic pr oper t y of met abolically act ive cell is-
(a) Low nucleo-cyt oplasmic r at io
(b) High nucleo-cyt oplasmic r at io
(c) High volume t o sur face ar ea r at io
(d) Small nucleus
Practice Test Paper–Cell Biology 39
18. Plasma membr ane t he funct ional as well as st r uct ur al r ole is played by-
(a) Pr ot eins (b) Lipids
(c) Cholest er ol (d) Oligosacchar ides
19. Lipid nat ur e of plasma membr ane can be dest r oyed by-
(a) Hexane (b) Benzene
(c) Chlor ofor m (d) NaOH
20. The plasma membr ane of int est ine is highly folded int o micr ovilli. The main funct ion
of Micr ovilli is-
(a) To Secr et e digest ive enzymes (b) To help in blood cir culat ion
(c) To incr ease it s absor pt ive sur face (d) For ageing of wor n out cells
21. The st r uct ur e for med wher e t wo adjacent membr ane ar e t hickened wit h disc shaped
adhesive mat er ial in bet ween and t onofibr ils r adiat ing out fr om adhesive r egion is-
(a) Gap junct ion (b) Tight junct ions
(c) Desmosomes (d) Plasmodesmat a
22. The out er par t of cyt oplasm is usually t er med as-
(a) Pla sma sol (b) Pla sma gel
(c) Nucleoplasm (d) Pr ot oplasm
23. Endoplasmic r et iculum or iginat es fr om-
(a) Nuclues (b) Nucleulous
(c) Golgi Complex (d) Plasma membr ane.
24. The endoplasmic r et iculum which const it ut e 50 % of cell is absent in-
(a) Ova (b) Embr yonic cells
(c) Mat ur e er yt hr ocyt es (d) All of t he above
25. Ribosome ar e at t ached t o endoplasmic r et iculum t hr ough glycopr ot ein known as
Ribophor in I & II. The Subunit of r ibosome which get at t ached t o ER is-
(a) P sit e (b) A sit e
(c) Lar ge subunit (d) Small subunit
26. How you can separ at e Gr am + ve bact er ia fr om Gr am –ve bact er ia-
(a) Pr esence of Techoic Acid (b) Absence of per iplasmic Space
(c) Exot oxin Pr oduced (d) All of t he above
27. Lysosomes ar e polymor phous or ganelles enclosed by a single membr ane. They cont ain
vast ar r ay of hydr olyt ic enzymes which can digest any for eign mat er ial except -
(a) Cellulose (b) St ar ch
(c) Glycogen (d) Lipids
28. Spect r in of er yt hr ocyt es and cyt ochr ome c of mit ochondr ia, which can be easily
dissociat ed by high ionic st r engt h and met al ion chelat ing agent ar e example of-
(a) Ext r insic Pr ot ein (b) Int r insic pr ot ein
(c) Tunnel Pr ot ein (d) Cyt oplasmic Pr ot ein
40 CSIR-NET Life Sciences
29. Fat s, St er ol and det oxificat ion ar e found abundant in-
(a) Adipose cells (b) Muscle cells
(c) Liver cells (d) All of above
30. RER is found abundant ly in goblet cells, pancr eat ic cells and liver cells is mainly
engaged in-
(a) Glycosylat ion of pr ot ein
(b) Folding and Secondar y St r uct ur e for mat ion
(c) Pr oduct ion of Secr et or y & cyt osolic pr ot ein
(d) Pr oduct ion and Excr et ion of pr ot ein
31. Micr osomes ar e not found in cell in nat ur al condit ion. They ar e-
(a) Pr esent only in cer t ain bact er ia
(b) Br oken pieces of ER dur ing cent r ifugat ion
(c) Br oken pieces of golgi dur ing cent r ifugat ion
(d) Pr esent in cer t ain fungi
32. Among t he following which is not pr esent in smaller subunit of r ibosome-
(a) Pept idyl t r ansfer ase (b) Binding sit e for t RNA
(c) A Sit e (d) P sit e
33. Polyr ibosome ar e seen in-
(a) Bact er ia (b) Fungi
(c) Angiosper ms (d) Mammals
34. r -RNA or iginat es fr om-
(a) Nucleus (b) Nucleolous
(c) Cyt oplasm (d) ER
35. Which or ganelle is pr esent ing zone of exclusion and have definit e polar it y-
(a) Golgi (b) ER
(c) Nucleus (d) Ribosome
36. Lysosomes ar e abundant in-
(a) WBC and ost eoblast s (b) RBC and Spleen
(c) Liver and Spleen (d) WBC and Spleen
37. Lysosome membr ane is st r engt hened by cor t isol, cor t isone, ant ihist amine, hepar in,
chlor oquinone and cholest er ol but becomes fr agile-
(a) Low bile salt s and ener gy r adiat ions
(b) In absence of oxygen
(c) Low Vit amin A & E
(d) Low Pr ogest r one and est r ogen
38. Fr uit r ot t ing can be checked by slowing down t he act ion of enzyme polygalact our onose
of t he or ganelle-
(a) Golgi (b) Lysosomes
(c) Glyoxysome (d) Per oxisome
Practice Test Paper–Cell Biology 41
39. In pr okar yot es wher e t he mit ochondr ia is absent , t he sit e of oxidat ive phosphor ylat ion
and elect r on t r anspor t chain including dehydr ogenases is-
(a) Mesosomes (b) Endosomes
(c) Plasma membr ane (d) Micr osomes
40. Wat er soluble phycobillin pigment occur in-
(a) BGA and Gr een algae (b) BGA and Red algae
(c) Gr een algae and Red algae (d) Gr een algae and Br own algae
41. Phot osynt het ic pigment s ar e locat ed in membr ane on specific ar eas called as-
(a) Oxysomes (b) Qua nt osomes
(c) Phot osyst em (d) Ant enna molecules
42. Mi cr ot u bu l es a r e 25 n m t h i ck , 15n m cor e for med of 13 h el i ca l l y a r r a n ged
pr ot ofilament s made up of-
(a) a t ubulin (b) b-t ubulin
(c) Myosin (d) bot h a & b
43. Int er mediat e filament s ar e made up of-
(a) Non-cont r act ile pr ot eins (b) b-t ubulin
(c) Myosin (d) Act in
44. In hexose m onophosphate shunt, the CO
2
molecules evolved is-
(a) Same as in glycolysis (b) Less t hen glycolysis
(c) Mor e t hen glycolysis (d) Much lesser t hen glycolysis
45. The elect r on donor dur ing nit r ogen fixat ion is-
(a) Wat er (b) Fer r ocynide
(c) Fer odoxin (d) CO
2
46. The chr omat in is made up of r epit at ive unit s known-
(a) Chr omosomes (b) Chr omonemat a
(c) Nucleosomes (d) Nucleot ides
47. Exocyt osis and endocyt osis is absent in-
(a) Amoeba (b) Euglena
(c) Mycoplasma (d) Algae
48. Cyt ochr ome oxidase and cyt ochr ome c deficiency in mit ochondr ia causes-
(a) Menke’s disease
(b) Kear ns-says syndr ome and Menke’s disease
(c) Kear ns-says syndr ome
(d) Leber ’s opt ic neur opat hy
49. Phot ophosophor ylat ion occur s in-
(a) Plast ids (b) Mit ochondr ia
(c) Cyt oplasm (d) Cell membr ane
42 CSIR-NET Life Sciences
50. Which of t he following is cor r ect wit h r egar d t o aneuploidy?
(a) inver sion
(b) 2n + 1
(c) All aneuploid individuals die befor e bir t h.
(d) 4n
51. I f a fr a gment of a chr omosome br ea ks off a nd t hen r ea t t a ches t o t he or igina l
chr omosome but in t he r ever se dir ect ion, t he r esult ing chr omosomal abnor malit y is
called
(a) a delet ion. (b) an inver sion.
(c) a t r anslocat ion. (d) a nondisjunct ion.
52. Why ar e individuals wit h an ext r a chr omosome 21, which causes Down syndr ome,
mor e numer ous t han individuals wit h an ext r a chr omosome 3 or chr omosome 16?
(a) Ther e ar e pr obably mor e genes on chr omosome 21 t han on t he ot her s.
(b) Chr omosome 21 is a sex chr omosome and 3 and 16 ar e not .
(c) Down syndr ome is not mor e common, just mor e ser ious.
(d) Ext r a copies of t he ot her chr omosomes ar e pr obably fat al.
53. Humans have 23 pair s of chr omosomes, while our closest r elat ives, chimpanzees,
have 24. Chr omosome st udies indicat e t hat at some point ear ly in human evolut ion,
t wo chr omosomes simult aneously br oke int o a lar ge por t ion and a small por t ion. The
lar ge par t s combined t o for m a lar ge chr omosome, and t he small par t s combined t o
for m a much smaller chr omosome (which was subsequent ly lost ). This impor t ant
chr omosomal change could best be descr ibed as
(a) nondisjunct ion followed by delet ion.
(b) t r anslocat ion followed by delet ion.
(c) duplicat ion followed by delet ion.
(d) t r anslocat ion followed by inver sion.
54. Each cell in an individual wit h Down syndr ome cont ains ____ chr omosomes.
(a) 3 (b) 22
(c) 24 (d) 47
55. Disor der s involving unusual number s of sex chr omosomes show t hat maleness is
caused by t he
(a) pr esence of a n X chr omosome.
(b) pr esence of a Y chr omosome.
(c) a bsence of a n X chr omosome.
(d) a bsence of a Y chr omosome.
56. A par t icular allele can have differ ent effect s if it was inher it ed fr om a male r at her
t han a female. This phenomenon is known as
(a) ext r anuclear inher it ance. (b) aneuploidy.
(c) sex-linkage. (d) genome impr int ing.
Practice Test Paper–Cell Biology 43
57. Human mit ochondr ia
(a) ar e inher it ed as an X-linked t r ait .
(b) ar e all inher it ed fr om t he fat her .
(c) have linear DNA.
(d) ar e all inher it ed fr om t he mot her .
58. Bot h chlor oplast s and mit ochondr ia
(a) ar e found wit hin t he nucleus.
(b) have linear DNA.
(c) car r y ext r anuclear genes.
(d) ar e inher it ed fr om bot h par ent s.
59. Damaged DNA is excised by
(a) r est r ict ion enzymes. (b) helicase.
(c) pr imase. (d) DNA polymer ase.
60. Unlike pr okar yot ic DNA r eplicat ion, eukar yot ic DNA r eplicat ion
(a) is complet ed by DNA polymer ase.
(b) cannot be complet ed by DNA polymer ase.
(c) is semiconser vat ive.
(d) has a mult iple or igin.
61. Which of t he following is an example of a hydr ophobic mat er ial?
(a) Paper (b) Sugar
(c) Past a (d) Wax
62. We can be sur e t hat a mole of t able sugar and a mole of vit amin C ar e equal in t heir
(a) weight in dalt ons
(b) number of molecules
(c) volume
(d) number of at oms
63. Among t he following which is longest cell-
(a) Hemp (b) Ramie
(c) J ut e (d) Ner ve fibr e
64. The middle lamella of plant cells is made up of calcium magnesium pect at e. Pect ic
Acid is polymer of-
(a) a-1,4 D-Glucose (b) b-1,6-D Glucose
(c) a-1,4 D-Galact our onic acid (d) b-1,4 D-Galact our onic acid
65. Acid pr ecipit at ion has lower ed t he pH of a par t icular lake t o 4.0. What is t he hydr ogen
ion concent r at ion of t he lake
(a) 10
–4
M (b) 4.0 M
(c) 10
–10
M (d) 10
4
M
44 CSIR-NET Life Sciences
66. The per cent age amount of Int egr al pr ot ein of plasma membr ane is-
(a) 40 % (b) 50 %
(c) 60 % (d) 70 %
67. Oligosacchar ide ar e usely at t ached t o ext r insic phase of plasma membr ane by-
(a) Pr ot eins (b) Lipids
(c) Bot h a & b (d) Not specific
68. Maximum number of enzymes in a eukar yot ic cell is pr esent inside
(a) Cyt osol (b) Mit ocondr ia
(c) Lysosome (d) ER
69. Which of t he following t er m includes all ot her s in t he list ?
(a) Monosacchar ide (b) Car bohydr at e
(c) Disacchar ide (d) St ar ch
70. The st r uct ur al level of a pr ot ein least affect ed by a disr upt ion in hydr ogen bond is t he
(a) secondar y level (b) t er t iar y level
(c) pr imar y level (d) quat er nar y level
71. To conver t a nucleoside t o a nucleot ide, it would be necessar y t o:
(a) r emove t he pent ose fr om t he nucleoside (b) add phosphat e t o t he nucleoside
(c) r eplace pur ine wit h pyr imidine (d) r eplace r ibose wit h deoxyr ibose
72. Choose t he pair of t er ms t hat cor r ect ly complet es t his sent ence:
Nucleot ides a r e t o _______________ a s _______________ a r e t o pr ot eins.
(a) amino acids; polypept ides (b) genes; enzymes
(c) nucleic acids; amino acids (d) polymer s, polypept ides
73. Post t r anslat ion modificat ion of secr et ar y pr ot eins occur s in:
(a) RER (b) SER
(c) Mit ocondr ia (d) nucleus
74. Most cells cannot har ness heat in or der t o per for m wor k because
(a) heat is not a for m of ener gy
(b) Cells do not have much heat ; t hey ar e r elat ively cool
(c) heat denat ur es enzymes
(d) t emper at ur e is usually unifor m t hr oughout a cell
75. Choose t he pair of t er ms t hat cor r ect ly complet es t his sent ence; Cat abolism is t o
anabolism as ––––––––– is t o ––––––––––.
(a) exer gonic ; spont aneous (b) fr ee ener gy ; ent r opy
(c) exer gonic ; ender gonic (d) wor k ; ener gy
76. Accor ding t o t he fir st law of t her modynamics
(a) mat t er can be neit her cr eat ed nor dest r oyed.
(b) all pr ocesses incr ease t he or der of t he univer se.
(c) syst ems r ich in ener gy ar e int r insically st able
(d) ener gy is conser ved in all pr ocesses
Practice Test Paper–Cell Biology 45
77. How you can separ at e Gr am + ve bact er ia fr om Gr am –ve bact er ia-
(a) Pr esence of Techoic Acid (b) Absence of per iplasmic Space
(c) Exot oxin Pr oduced (d) All of t he above
78. The main Phagocyt ot ic cell in immune r esponse is-
(a) Neut r ophils (b) Basophils
(c) Monocyt es (d) Lymphocyt es
79. The phagocyt es ar e at t r act ed t owar d micr oor ganisms by-
(a) Chemot axis (b) Rheot axis
(c) Diapedesis (d) Thogmot axis
80. The cell wall of micr oor ganisms is coat ed wit h cer t ain plasma pr ot ein pr omot ing t he
at t achment of micr obe t o phagocyt es, only t hen t hey can be phagocyt osised. The coat
pr ot ein ar e called as-
(a) Globins (b) Opsonins
(c) Ovulbumins (d) Phagosonins
81. Aft er t he damage of body t issues, blood vessel is dilat ed wher e damage has occur r ed,
due t o which per meabilit y of blood vessel also incr eases. Vasodilat ion is caused by-
(a) Hist amine (b) Kinin
(c) Pr ost aglandin (d) All of above
82. The pr ocess of sneezing of phagocyt es bet ween t he endot helial cells of blood vessels
and r eaching t o damaged ar ea is known as-
(a) Mar ginat ion (b) Met ast asis
(c) Diapedesis (d) Angiobiosis
83. In humans int er fer on is pr oduced by leucocyt es, fibr oblast s in connect ive t issue and
lymphocyt es and ar e t er med as a-IFN, b-IFN and g-INF r esp. The int er fer ons ar e-
(a) Ant ibact er ial pr ot eins (b) Ant ivir al Pr ot ein
(c) Ant i cancer ous pr ot ein (d) Ant icancer pr ot ein
84. Dur ing embr yonic st age of human B-lymphocyt es ar e pr oduced in-
(a) Bone ma r r ow (b) Spleen
(c) Liver (d) Bur sa
85. Hapt ens ar e –
(a) Immunogenic ant igen
(b) Non Immunogenic Ant igen
(c) High molecular weight non immunogenic ant igen
(d) Low molecular weight immunogenic ant igen
86. Lysosomes ar e polymor phous or ganelles cont aining vast ar r ay of hydr olyt ic enzymes
which can digest any for eign mat er ial only at pH-
(a) 5 (b) 6
(c) 9 (d) 7
46 CSIR-NET Life Sciences
87. Number of ant igen funct ional binding sit e in human Immunoglobin-M ar e-
(a) 2 (b) 5
(c) 10 (d) 20
88. Number of a mino a cids in light a nd hea vy cha in of t ypica l immunoglobin a r e
r espect ively-
(a) 110, 220 (b) 220, 440
(c) 440, 880 (d) 880, 1760
89. Among t he following which is not essent ial pr oper t y of immunoglobin-
(a) Memor y (b) Specificit y
(c) Diver sit y (d) React ivit y
90. Spect r in of er yt hr ocyt es and cyt ochr ome c of mit ochondr ia, which can be easily
dissociat ed by high ionic st r engt h and met al ion chelat ing agent ar e example of-
(a) Ext r insic Pr ot ein (b) Int r insic pr ot ein
(c) Tunnel Pr ot ein (d) Cyt oplasmic Pr ot ein
91. Polyclonal ant ibodies ar e-
(a) Clones against single ant igen by many B-cells
(b) Clones against single ant igen by single B-cells
(c) Clones against many ant igen by single B-cells
(d) Clones against many ant igen by many B-cells
92. Lymphokines t hat r ecr uit t he macr ophages for Phagocyt osis ar e secr et ed by-
(a) T-cells (b) B-cells
(c) Complement syst em (d) MHC
93. Liposomes a r e-
(a) Lipid filled bags (b) Ar t ificial membr anes
(c) Liver Fat bodies (d) Eukar yot ic or ganelle
94. Which among t he following act as br idge bet ween cell mediat ed and humor al immunit y-
(a) T-cyt oxic cells (b) T-suppr essor cells
(c) B-cells (d) T-helper cells
95. In humans cell r ecognit ion molecules ar e-
(a) HLA (b) B-cells
(c) T-Cells (d) Immunoglobins
96. Cancer s cell ar e monoclonal, ar e char act er ized by uncont r olled gr owt h, invasion of
ot her t issues and disseminat ion t o ot her t issues. The phenomenon of invasion t o
ot her t issues is t er med as-
(a) Angiobiogenesis (b) Met ast asis
(c) Diapedesis (d) Tr ansfor mat ion
Practice Test Paper–Cell Biology 47
97. A major pr ot ease secr et ed by cancer cells act s on plasminogen and conver t s it int o
plasmin. Plasmin is pr ot eolyt ic enzyme t hat dissolves blood clot s and also r emoves
exposed pr ot ein gr oups at cell sur face. If t he plasminogen is r emoved for m t he medium,
t hen-
(a) The mor phology of cancer cells r et ur ns t o nor mal.
(b) The cancer cell will show mor e exponent ial gr owt h.
(c) Cancer cell will die
(d) No change will be seen
98. One of t he major higher molecular weight glycopr ot ein component which can be
easily isolat ed fr om nor mal cult ur ed fibr oblast by mild t r eat ment of ur ea, also occur
at “foot pr int s” t hat moving cult ur e cells leaves, is t ot ally absent in cancer ous cell is-
(a) Fibr onect in (b) Albumin
(c) Fer r it in (d) Tr ansfer in
99. The sex det er minat ion in dr ossophila is based on-
(a) X-Chr omosome (b) Y chr omosome
(c) Aut osome (d) Bot h a & c
100. Envir onment al cont r ol of sex det er minat ion is seen in-
(a) Melandr ium (b) Dr osophila
(c) Bonelia (d) Apes indica
Practice Test Paper-II
1. Which of t he following is not a funct ion of mit ot ic cell division in
(a) r epair of damaged or gans (b) pr oduct ion of gamet es
(c) asexual r epr oduct ion (d) gr owt h
2. Which of t he following would not be consider ed par t of a cell’s cyt oplasm?
(a) a r ibosome (b) t he nucleus
(c) a mit ochondr ion (d) a micr ot ubule
3. It is difficult t o obser ve individual chr omosomes wit h a light micr oscope dur ing
int er phase because:
(a) t hey have uncoiled t o for m long, t hin st r ands
(b) t hey leave t he nucleus and ar e disper sed t o ot her par t s of t he cell
(c) t he DNA has not been r eplicat ed yet
(d) t he spindle must move t hem t o t he met aphase plat e befor e t hey become
4. The maximum size of a cell is limit ed by
(a) it s need for enough sur face ar ea for exchange wit h it s envir onment .
(b) t he number of or ganelles t hat can be packed inside.
(c) t he mat er ials needed t o build it .
(d) t he amount of flexibilit y it needs t o be able t o move.
5. You would expect a cell wit h an ext ensive Golgi appar at us t o
(a) make a lot of ATP. (b) secr et e a lot of mat er ial.
(c) move act ively. (d) per for m phot osynt hesis.
6. Which of t he following cor r ect ly mat ches an or ganelle wit h it s funct ion?
(a) mit ochondr ion . . . phot osynt hesis
(b) nucleus . . . cellular r espir at ion
(c) r ibosome . . . manufact ur e of lipids
(d) cent r al vacuole . . . st or age
7. Mit ochondr ia and cholor plast s shar e sever al common feat ur es, for example,
(a) bot h ar e capable of semiaut onomous gr owt h and r epr oduct ion.
(b) neit her ar e component s of t he endomembr ane syst em.
(c) each cont ains a small amount of DNA
(d) all of t he above.
8. In muscle cells t he ____ is specialized for t he st or age and r elease of calcium.
(a) smoot h ER (b) t he Golgi appar at us
(c) cont r act ile vacuoles (d) r ough ER
Practice Test Paper–Cell Biology 49
9. Of t he following or ganelles, which gr oup is involved in manufact ur ing subst ances
needed by t he cell?
(a) lysosome, va cuole, r ibosome
(b) r ibosome, r ough ER, smoot h ER
(c) vacuole, r ough ER, smoot h ER
(d) smoot h ER, r ibosome, vacuole
10. The int er nal skelet on of a cell is composed of
(a) micr ot ubules, int er mediat e filament s, and micr ofilament s.
(b) cellulose and int er mediat e filament s.
(c) cellulose, micr ot ubules, and cent r ioles.
(d) micr ofilament s.
11. Cells will usually divide if t hey r eceive t he pr oper signal at a
(a) M (b) S
(c) G2 (d) G1
12. Dye inject ed int o a plant cell might be able t o ent er an adjacent cell t hr ough a
(a) t ight junct ion. (b) micr ot ubule.
(c) desmosome. (d) plasmodesma.
13. A r esear cher made an int er est ing obser vat ion about a pr ot ein made by t he r ough ER
and event ually used t o build a cell’s plasma membr ane. The pr ot ein in t he membr ane
was act ually slight ly differ ent fr om t he pr ot ein made in t he ER. The pr ot ein was
pr obably changed in t he
(a) Golgi appar at us. (b) smoot h ER.
(c) mit ochondr ion. (d) nucleus.
14. When elongat ed, t ube-shaped cells fr om t he lining of t he int est ine ar e t r eat ed wit h a
cer t ain chemical, t he cells sag and become r ound blobs. The int er nal st r uct ur es
disr upt ed by t his chemical ar e pr obably
(a) cell junct ions. (b) micr ot ubules.
(c) r ough ER. (d) dynein.
15. The elect r on micr oscope has been par t icular ly useful in st udying bact er ia, because
(a) elect r ons can penet r at e t ough bact er ial cell walls.
(b) bact er ia ar e so small.
(c) bact er ia move so quickly t hey ar e har d t o phot ogr aph.
(d) wit h few or ganelles pr esent , bact er ia ar e dist inguished by differ ences in individual
ma cr omolecules.
16. Cell fr act ionat ion is t he most appr opr iat e pr ocedur e for pr epar ing ____ for st udy.
(a) isolat ed cells which ar e nor mally found t ight ly at t ached t o neighbour ing cells
(b) cells wit hout a funct ional cyt oskelet on
(c) isolat ed or ganelles
(d) t he basic macr omolecules
50 CSIR-NET Life Sciences
17. Which of t he following clues would t ell you whet her a cell is pr okar yot ic or eukar yot ic?
(a) t he pr esence or absence of a r igid cell wall
(b) whet her or not t he cell is par t it ioned by int er nal membr anes
(c) t he pr esence or absence of r ibosomes
(d) whet her or not t he cell car r ies out cellular met abolism
18. Seema would like t o film t he movement of chr omosomes dur ing cell division. Her
best choice for a micr oscope would be a
(a) t r ansmission elect r on micr oscope, because of it s magnifying power .
(b) scanning elect r on micr oscope, because t he specimen is alive.
(c) t r ansmission elect r on micr oscope, because of it s gr eat r esolving power .
(d) light micr oscope, because t he specimen is alive.
19. A plant cell was gr own in a t est t ube cont aining r adioact ive nucleot ides, t he par t s
fr om which DNA is built . Lat er examinat ion of t he cell showed t he r adioact ivit y t o be
concent r at ed in t he
(a) r ough ER. (b) per oxisome.
(c) nucleus. (d) cent r al vacuole.
20. When isolat ed liver cells ar e combined wit h nonpolar t oxins init ial pr ocessing in t he
___ incr eases t he solubilit y of t hese compounds as an init ial st ep in t heir excr et ion.
(a) smoot h ER (b) Golgi appar at us
(c) mit ochondr ion (d) r ough ER
21. Discont inuous segment s of DNA ar e joined by:
(a) t elomer ase (b) DNA ligase
(c) exonuclease (d) DNA polymer ase
22. The concent r at ion of calcium in a cell is 0.3%. The concent r at ion of calcium in t he
sur r ounding fluid is 0.1%. How could t he cell obt ain mor e calcium?
(a) passive t r anspor t (b) diffusion
(c) act ive t r anspor t (d) osmosis
23. Phospholipid molecules in a membr ane ar e ar r anged wit h t heir ____ on t he ext er ior
and t heir ____ on t he int er ior .
(a) hydr ophobic heads . . . hydr ophilic t ails
(b) hydr ophilic heads . . . hydr ophobic t ails
(c) nonpolar heads . . . polar t ails
(d) hydr ophobic t ails . . . hydr ophilic heads
24. Sist er chr omat ids:
(a) ar e cr eat ed when DNA is r eplicat ed
(b) ar e separ at ed dur ing mit osis
(c) ar e at t ached at t he cent r omer e pr ior t o division
(d) all of t he above
Practice Test Paper–Cell Biology 51
25. Which of t he following cor r ect ly mat ches a phase of t he cell cycle
(a) S, immediat ely pr ecedes cell division (b) M, duplicat ion of DNA
(c) G1, immediat ely follows cell division (d) G2, cell division
26. The kinet ochor es:
(a) ar e t he pr imar y cent r omer e st r uct ur es t hat maint ain t he at t achment
(b) at t ach t o t he r ing of act in along t he cyt oplasmic sur face of t he plasma membr ane
(c) move along spindle micr ot ubules dur ing anaphase, dr agging
(d) ar e locat ed at t he cent er of t he cent r omer e t heir funct ion is t o pr ovide at t achment
sit e t o spindle fibr e
27. A biochemist measur ed t he amount of DNA t o be doubled dur ing t he cells in gr owing
st ages
(a) bet ween pr ophase and anaphase
(b) bet ween anaphase and t elophase
(c) dur ing t he M phase of t he cell cycle
(d) bet ween t he G1 and G2 phases
28. DNA damaged by sunlight :
(a) has under gone depur inat ion (b) has lost it s phosphat e gr oups
(c) has for med pyr imidine dimer s (d) has lost it s hydr ogen bonds
29. Which of t he following funct ional pr ocesses r esult s fr om t he pr esence of pr ot ein wit hin
t he plasma membr ane?
(a) enzymat ic act ivit y (b) signal t r ansduct ion
(c) int er cellular joining (d) all of t he above
30. Select t he cor r ect st at ement concer ning membr ane car bohydr at e.
(a) Car bohydr at es ar e only found associat ed wit h t he membr anes of pr okar yot ic
cells.
(b) Glucose is t he most abundant membr ane car bohydr at e.
(c) Cell membr a nes consist of pr ot ein a nd phospholipid; ca r bohydr a t e is not a
membr ane component .
(d) membr ane car bohydr at es funct ion pr imar ily in cell-cell r ecognit ion.
31. Imagine t wo solut ions separ at ed by a select ively per meable membr ane which allows
wat er t o pass, but not sucr ose or glucose. The membr ane separ at es a 0.2 M sucr ose
solut ion fr om a 0.2 glucose solut ion. Wit h t ime how will t he solut ions change?
(a) Not hing happens because t he t wo solut ions ar e isot onic t o one anot her .
(b) Wat er ent er s t he sucr ose solut ion because t he sucr ose molecule is a disacchar ide,
and is lar ger t han t he monosacchar ide glucose.
(c) Wat er leaves t he sucr ose solut ion because t he sucr ose molecule is a disacchar ide,
and is lar ger t han t he monosacchar ide glucose.
(d) The sucr ose solut ion is hyper t onic and will gain wat er because t he t ot al mass of
sucr ose is gr eat er t han t hat of glucose.
52 CSIR-NET Life Sciences
32. Dur ing cyt okinesis in plant s:
(a) a bundle of act in micr ofilament s called t he cont r act ile r ing, pinch t he cell in half
(b) small vesicles, dir ect ed by t he phr agmoplast , move t o t he spindle
(c) a cleavage fur r ow encir cles t he cell
(d) cyt oplasmic division is called cleavage
33. Pr ogr ession t hr ough t he eukar yot ic cell cycle is r egulat ed by:
(a) micr ot ubules (b) t he p53 gene
(c) cyclin-dependent kinases (d) DNA ligase
34. The r esult of t he oper at ion of an elect r ogenic pump would be
(a) an elect r ochemical gr adient on t he cell membr ane
(b) a r est ing pot ent ial acr oss t he membr ane
(c) plasmolysis
(d) a cell wit h a posit ively char ged int er ior
35. A plant cell is placed in a solut ion whose solut e concent r at ion is t wice as gr eat as t he
concent r at ion of t he cell cyt oplasm. The cell membr ane is select ively per meable,
allowing wat er but not t he solut es t o pass t hr ough. What will happen t o t he cell?
(a) No change will occur because it is a plant cell.
(b) The cell will shr ivel because of osmosis.
(c) The cell will swell because of osmosis.
(d) The cell will shr ivel because of act ive t r anspor t of wat er .
36. One consequence of t he sidedness of t he plasma membr ane is t hat
(a) molecules t hat begin on t he inside face of t he ER end up on t he inside face of t he
plasma membr ane
(b) t he asymmet r ical dist r ibut ion for membr ane pr ot eins, lipids, and car bohydr at e
must be det er mined when t he membr ane is fir st const r uct ed
(c) some pr ot eins on t he cyt oplasmic side of t he membr ane ar e at t ached t o t he
cyt oskelet on
(d) t he inside of an ER vesicle is t opogr aphical equivalent t o t he ext r a cellular sur face
of t he plasma membr ane
37. The p53 gene:
(a) is t he most fr equent ly mut at ed gene in human cancer
(b) can lead t o cell cycle ar r est at t he G1 checkpoint
(c) can t r igger apopt osis.
(d) all of t he above
38. Dur ing t he Ras pat hway:
(a) cyt oplasmic pr ot ein kinases ar e act ivat ed
(b) t he gr owt h fact or r ecept or is dephosphor ylat ed
(c) gr owt h fact or s bind t o r ecept or s in t he cyt oplasm
(d) leads t o t he pr oduct ion of t r anslat ion fact or s
Practice Test Paper–Cell Biology 53
39. If a DNA molecule has a deaminat ed base, it will be r epair ed by:
(a) by excision r epair pat hways
(b) wit h t he help of r epair endonucleases
(c) by base excision r epair
(d) wit h t he help of DNA glycosylases
40. A benign t umor is one in which t he cancer ous cells:
(a) have an unusual number of chr omosomes
(b) can divide indefinit ely if an adequat e supply of nut r ient s is available
(c) migr at e fr om t he init ial sit e of t r ansfor mat ion t o ot her or gans or
(d) r emain confined t o t heir or iginal sit e
41. A DNA molecule is a polymer made of subunit s called
(a) bases. (b) amino acids.
(c) nucleot ides. (d) nucleic acids.
42. A human somat ic cell cont ains _____ chr omosomes.
(a) 23 (b) 47
(c) 2n (d) 46
43. Which of t he following is a nor mal human female?
(a) X Y (b) X X Y
(c) X X (d) X
44. A kar yot ype would be least likely t o show which of t he following?
(a) an ext r a chr omosome
(b) par t of a chr omosome duplicat ed
(d) a missing chr omosome
(d) par t of a chr omosome t ur ned ar ound
45. The diploid st age of a plant t hat exhibit s an alt er nat ion of gener at ion is t he
(a) ant her idium. (b) gamet ophyt e.
(c) spor e. (d) spor ophyt e.
46. DNA r eplicat ion occur s
(a) whenever a cell makes pr ot ein.
(b) t o r epair gene damage caused by mut at ion.
(c) befor e a cell divides.
(d) whenever a cell needs RNA.
47. Mit osis and cyt okinesis r esult in t he for mat ion of ______; meiosis and cyt okinesis
r esult in t he for mat ion of ________.
(a) 4 diploid cells; 4 haploid cells
(b) 2 diploid cells; 2 haploid cells
(c) 2 diploid cells; 4 haploid cells
(d) 2 diploid cells; 2 diploid cells.
54 CSIR-NET Life Sciences
48. Which of t he following occur s dur ing meiosis but not dur ing mit osis?
(a) Chr omosomes align at t he met aphase plat e.
(b) Chr omosomes condense.
(c) Chr omosomes migr at e t o opposit e poles.
(d) synapsis
49. At t he end of t elophase I and cyt okinesis t her e ar e
(a) 4 haploid cells. (b) 2 diploid cells.
(c) 1 haploid ovum and 3 polar bodies. (d) 2 haploid cells.
50. Synapsis occur s dur ing
(a) anaphase I. (b) pr ophase I.
(c) cyt okinesis. (d) pr ophase II.
51. Dur ing anaphase II
(a) homologues separ at e and migr at e t owar ds opposit e poles.
(b) sist er chr omat ids separ at e and migr at e t owar ds opposit e poles.
(c) nuclei r efor m.
(d) chr omosomes line up in a single file.
52. Dur ing anaphase I
(a) homologues separ at e and migr at e t owar ds opposit e poles.
(b) sist er chr omat ids separ at e and migr at e t owar ds opposit e poles.
(c) nuclei r efor m.
(d) chr omosomes line up in a single file.
53. Cyt okinesis is t he
(a) exchange of homologous r egions of nonsist er chr omat ids.
(b) for mat ion of t et r ads.
(c) independent assor t ment of chr omosomes.
(d) division of one cell int o t wo
54. Cent r ioles separ at e dur ing
(a) cyt okinesis.
(b) pr ophase I and pr ophase II
(c) met aphase I and met aphase II.
(d) anaphase I and anaphase II.
55. Cr ossing over occur s dur ing
(a) cyt okinesis. (b) met aphase I.
(c) pr ophase II. (d) pr ophase I
56. Regions wher e nonsist er chr omat ids cr oss ar e called
(a) inver sions. (b) homologues.
(c) t et r ads (d) chiasmat a.
Practice Test Paper–Cell Biology 55
57. In humans, t he haploid number of chr omosomes is 23. Independent assor t ment has
t he possibilit y of pr oducing _____ differ ent gamet es.
(a) 23
2
(b) 8 million
(c) 22
3
(d) 24
58. Cr ossing over is
(a) t he movement of genet ic ma t er ia l fr om one chr omosome t o a nonhomologous
chr omosome.
(b) independent assor t ment of chr omosomes.
(c) t he for mat ion of chiasmat a.
(d) t he exchange of homologous por t ions of nonsist er chr omat ids.
59. Var iat ion occur s when chr omosomes ar e shuffled in _____ and fer t ilizat ion.
(a) mit osis (b) genet ic dr ift
(c) meiosis (d) mut at ion
60. Her it able var iat ion is r equir ed for
(a) meiosis. (b) mit osis.
(c) evolut ion. (d) asexual r epr oduct ion.
61. An ant igen is
(a) a pr ot ein molecule t hat helps defend t he body against disease.
(b) a t ype of whit e blood cell.
(c) an invading vir us or bact er ium.
(d) a for eign molecule t hat evokes a specific r esponse by a lymphocyt e.
62. How do memor y cells differ fr om effect or cells?
(a) Memor y cells a r e mor e numer ous.
(b) Memor y cells ar e r esponsible for t he pr imar y immune r esponse.
(c) Memor y cells at t ack invader s; effect or cells do not .
(d) Memor y cells live longer .
63. Following t issue damage or t he ent r y of micr oor ganisms, an inflammat or y r esponse
may be init iat ed by
(a) t he accumulat ion of phagocyt es in an injur ed ar ea.
(b) t he r elease of int er fer on by infect ed cells.
(c) an incr eased blood flow in an infect ed or injur ed ar ea.
(d) t he r elease of chemicals such as hist amine by damaged cells.
64. Which of t he following could be consider ed a nonspecific defense?
(a) Int act skin cr eat es a bar r ier t hat cannot nor mally be penet r at ed by bact er ia or
vir uses.
(b) Secr et ions fr om sebaceous and sweat glands give t he skin an acidic pH t hat
pr event s bact er ial colonizat ion.
(c) Tear s, saliva, and mucous secr et ions cont ain lysozome, an enzyme t hat digest s
t he bact er ial cell wall.
(d) all of t he above
56 CSIR-NET Life Sciences
65. Dur ing a secondar y immune r esponse
(a) select ed B gener at e ant ibody-pr oducing effect or B cells called plasma cells.
(b) t he st r icken individual may become ill.
(c) about 10 t o 17 days ar e r equir ed fr om exposur e t o maximum effect or r esponse.
(d) t he gener at ion of effect or cells begins wit h memor y cells pr oduced dur ing t he
pr imar y immune r esponse.
66. Most individuals infect ed wit h HIV
(a) can live for 15 t o 20 year s.
(b) never develop AIDS.
(c) die fr om aut oimmune r eact ions.
(d) die fr om ot her infect ions or cancer .
67. Tissues ar e t yped befor e an or gan t r ansplant t o make sur e t hat t he _____ of donor
and r ecipient mat ch as closely as possible.
(a) T cells
(b) ant ibodies
(c) MHC (major hist ocompat ibilit y complex) pr ot eins
(d) hist amines
68. A vaccine cont ains
(a) whit e blood cells t hat fight infect ion.
(b) ant ibodies t hat r ecognize invading micr obes.
(c) inact ivat ed disease-causing micr obes.
(d) a hor mone t hat boost s immunit y.
69. When you ar e immune t o a disease,
(a) ant ibodies against t he disease ar e const ant ly cir culat ing in your blood.
(b) cer t ain lymphocyt es ar e able t o make t he pr oper ant ibodies quickly.
(c) your nonspecific defenses ar e st r engt hened.
(d) B cells ar e st imulat ed t o quickly engulf invader s.
70. In a ser ies of immune syst em exper iment s, t he t hymus glands wer e r emoved fr om
baby mice. Which of t he following would you pr edict as a likely r esult ?
(a) The mice suffer ed fr om numer ous aller gies.
(b) The mice never developed cancer ous t umor s.
(c) The mice suffer ed fr om aut oimmune diseases.
(d) The mice r eadily accept ed t issue t r ansplant s.
71. The body pr oduces ant ibodies complement ar y t o for eign ant igens. The pr ocess by
which t he body comes up wit h t he cor r ect ant ibodies t o a given disease is most like
(a) going t o a t ailor and having a suit made t o fit you.
(b) or der ing t he lunch special at a r est aur ant wit hout looking at t he menu.
(c) going t o a shoe st or e and t r ying on shoes unt il you find a pair t hat fit s.
(d) select ing a lot t er y pr ize-winner by means of a r andom dr awing.
Practice Test Paper–Cell Biology 57
72. The ant igen-binding sit es of an ant ibody molecule ar e for med fr om t he molecule’s
var iable r egions. Why ar e t hese r egions called var iable?
(a) They can change t heir shapes on command t o fit differ ent ant igens.
(b) They change t heir shapes when t hey bind t o an ant igen.
(c) They can be differ ent shapes on differ ent ant ibody molecules.
(d) Their sizes var y consider ably fr om one ant ibody t o anot her .
73. The biggest differ ence bet ween cell-mediat ed immunit y and humor al immunit y is
(a) how long t heir pr ot ect ion last s.
(b) whet her a subsequent secondar y immune r esponse can occur .
(c) whet her clonal select ion occur s.
(d) how t hey r espond t o and dispose of invader s.
74. Vir uses and bact er ia in body fluids ar e at t acked by
(a) ant ibodies fr om B cells. (b) cyt ot oxic T cells.
(c) complement pr ot eins. (d) helper T cells.
75. Wha t do t he a nt ibodies secr et ed by pla sma cells (t he effect or cells of humor a l
immunit y) do t o at t ack t heir t ar get s?
(a) act ivat e complement t o punch holes in t hem
(b) clump cells t oget her so t hat phagocyt es can ingest t hem
(c) cause ant igen molecules t o set t le out of solut ion
(d) all of t he above
76. Tissue macr ophages
(a) begin t heir lives as neut r ophils.
(b) have shor t life spans because t hey self-dest r uct aft er engulfing for eign invader s.
(c) or iginat e fr om monocyt es t hat leave t he cir culat ion and ent er t he t issues.
(d) ar e most effect ive against par asit es.
77. The idea behind vaccinat ion is t o induce _____ wit hout t he vaccinat ed individual
having t o get sick.
(a) passive immunit y
(b) t he pr imar y immune r esponse
(c) anaphylact ic shock
(d) nonspecific defenses
78. A gr oup of r esear cher s have t est ed many chemicals and found sever al t hat have
pot ent ial for use in modifying t he act ion of t he immune syst em. Which of t he following
would seem t o have t he most pr omise as a dr ug for inhibit ing t r ansplant r eject ion?
(a) Compound A13: act s like hist amine
(b) Compound Q6: suppr esses cyt ot oxic T cells
(c) Compound N98: a pot ent aller gen
(d) Compound M31: st imulat es helper T cells
58 CSIR-NET Life Sciences
79. Collagens:
(a) ar e high in glycine, hydr oxylysine and hydr oxypr oline
(b) ar e t he most abundant pr ot eins in ver t ebr at es
(c) ar e t he most abundant component of t he ext r acellular mat r ix in animals
(d) all of t he above
80. The ECM of animals cells consist s of what t hr ee classes of molecules?
(a) pr ot ein-polysacchar ides, st r uct ur al pr ot eins, micr ot ubules
(b) st r uct ur al pr ot eins, lipid bilayer s, adhesive glycopr ot eins.
(c) lipopr ot eins, polysacchar ides, and adhesive glycopr ot eins.
(d) st r uct ur al pr ot eins, pr ot ein-polysacchar ides, adhesive glycopr ot eins
81. Pr ocollagen consist s of _______ and is for med in t he lumen of t he:
(a) 2 b sheet s, lysosomes
(b) 3 b sheet s, E R
(c) 3 a chains, Golgi
(d) 3 a chains, E R
82. The flexibilit y of t he lungs is due mainly t o:
(a) laminins (c) elast ins
(b) collagens (d) fibr onect ins
83. A differ ence bet ween collagen and elast in is t hat :
(a) elast in has hydr oxylysines and collagen does not
(b) collagen does not have hydr oxylat ed pr oline r esidues
(c) t her e ar e over 15 t ypes of elast ins and less t han 5 t ypes of collagens
(d) elast in molecules ar e cr osslinked by covalent bonds bet ween lysine r esidues,
and collagen is not
84. Which is t he cor r ect or der fr om least t o most complex?
(a) Glucur onat e, hyalur onat e, pr ot eoglycans, GAG
(b) Hyalur onat e, glucur onat e, GAG chains, pr ot eoglycans
(c) GAG chains, hyalur onat e, glucur onat e, pr ot eoglycans
(d) Glucur onat e, hyalur onat e, GAG chains, pr ot eoglycans
85. Two of t he most common adhesive glycopr ot eins in t he ECM ar e:
(a) laminins and elast ins
(b) collagens and elast ins
(c) fibr onect ins and laminins
(d) collagens and fibr onect ins
86. Fibr onect ins have an effect on:
(a) blood clot t ing (b) cell shape
(c) cell movement (d) all of t he above
Practice Test Paper–Cell Biology 59
87. The most abundant glycopr ot eins in t he basal lamina ar e ______
(a) collagens (b) ent act in
(c) laminins (d) fibr onect ins
88. The most abundant pr ot ein in human body is-
(a) Collagen (b) Fibr onect in
(c) Globin (d) albumin
89. Roles of t he glycocalyx include:
(a) cell adhesion (b) cell r ecognit ion
(c) cr eat ion of per meabilit y bar r ier s (d) all of t he above
90. A r ed blood cell has just been t ar get ed for dest r uct ion. This cell:
(a) has an incr ease in sialic acid gr oups.
(b) has a decr ease in sialic acid gr oups
(c) has exposed galact ose r esidues
(d) B and C
91. The t hr ee most common cell junct ions in animals ar e:
(a) adhesive junct ions, t ight junct ions, gap junct ions
(b) adhesive junct ions, t ight junct ions, plasmodesmat a
(c) t ight junct ions, plasmodesmat a, adhesive junct ions
(d) gap junct ions, t ight junct ions, plasmodesmat a
92. The r eason ur ine does not seep out of t he bladder is due t o t he r ole of:
(a) adher ens junct ions (b) hemidesmosomes
(c) gap junct ions (d) t ight junct ions
93. Desmosomes and hemidesmosomes ar e anchor ed in t he cyt oplasm by:
(a) desmocollins (c) act in fiber s
(b) collagen (d) t onofilament s
94. Gap junct ions consist of channels for med by:
(a) hydr ophilic, connexons (b) hydr ophobic, cadher ins
(c) hydr ophobic, connexons (d) hydr ophilic, cadher ins
95. A plant cell wall includes all of t he following EXCEPT:
(a) hemicellulose (b) ext ensins
(c) cellulose (d) collagen
96. Which of t he following would be most likely t o lead t o cancer ?
(a) hyper act ivit y of bot h a pr ot o-oncogene and a t umor -suppr essor
(b) amplificat ion of a pr ot o-oncogene and t he failur e of a t umor -suppr essor gene
(c) hyper act ivit y of bit h a pr ot o-oncogene and a t umor suppr essor gene
(d) failur e of a pr ot o-oncogene t o pr oduce a pr ot ein and amplificat ion of t umor
suppr essor gene
60 CSIR-NET Life Sciences
97. The char act er ist ic st r engt h of woody t issues is due t o:
(a) t he secondar y cell wall (b) hemicellulose
(c) t he middle lamina (d) t he pr imar y cell wall
98. The r ule is der ived fr om t he ER and is cont inuous wit h t he ER of t wo adjacent plant
cells.
(a) middle lamina (b) annulus
(c) plasmodesma (d) desmot ubule
99. One can est imat e t he lengt h of M phase by knowing:
(a) t he S phase (c) t he age of t he cell
(b) t he mit ot ic index (d) t he gener at ion t ime
100. What is t r ue of pr ot o-oncogenes?
(a) cells pr oduce pr ot o-oncogenes as a by-pr oduct of mit osis
(b) pr ot o-oncogenes ar e necessar y for nor mal cont r ol of cell division
(c) pr ot o-oncogenes ar e genet ic junk t hat has not yet been eliminat ed by nat ur al
select ion
(d) pr ot o-oncogenes ar e unavoidable envir onment al car cinogens.
2
Biochemistry
BASIC CHEMISTRY
I. Properties of water
A. Wat er is polar
B. Effect on non-covalent bonds
1. elect r ost at ic A
+
......... B

2. hydr ogen bonds –A–H.......B

(A,B=N,O,S)
3. hydr ophobic bonds (exclusion by wat er )
4. van der Waals for ces opt imal dist ance (Vander wall cont act dist ance)
C. Wat er ionizat ion
1. H
2
O < > H
+
+ OH

2. Keq =
] 5 . 55 [
] OH ][ H [
O H
] OH ][ H [
2
− + − +
=
Kw = Keq × 55.5 = [H
+
] [OH

] = 1 × 10
–14
at 25
o
C
3. Pur e wat er : [H
+
] = [OH

] = 1 × 10
–7
II. Definition of pH and pOH
A. Gener al: p“X” = log (1/“X”) = –log“X”
B. pH = –log[H
+
]
C. pOH = –log[OH

]
D. pKw = –log Kw = pH + pOH = 14
E. pKa = –log Ka
F. A t en-fold change in [H
+
] alt er s t he pH by 1.0 unit .
62 CSIR-NET Life Sciences
III. Buffers
1. A buffer is a mixt ur e of a weak acid and t he salt of t hat acid.
2. The pr act ical buffer zone is pKa n
~
1 pH.
3. The Hender son-Ha ssa lba lch equa t ion is used for ca lcula t ions involving buffer s.
] HA [
] A [
log pKa pH

+ =
4. pH of a buffer is t heor et ically independent of dilut ion (depends only on t he r at ion of
A

/HA and pKa).
IV. Solving pH problems
A. St r ong acids (complet ely dissociat ed)
[H
+
] = nor malit y of acid pH = –log[H
+
]
B. St r ong bases (complet ely dissociat ed)
[OH

] = nor malit y of base pOH = –log[OH

] pH + pOH = 14
C. Weak acids: t wo kinds of pr oblems
1. % ionizat ion x molar it y = [H
+
]
2.
] HA [
x
~
x ] HA [
x
] HA [
] A ][ H [
Ka
2 2

− +
= =
x = [H
+
] = {Ka x [HA]}

V. Atoms
• At oms ar e like a planet ar y syst em wit h negat ive elect r ons cir cling ar ound a posit ively
char ged nucleus
• In elect r icit y opposit e char ges at t r act one anot her ; like char ges r epel
• All at oms ar e for med by t he at t r act ion of opposit ely char ged par t icles
(elect r ons & pr ot ons)
• Nucleus: has pr ot ons wit h (+) char ge and neut r ons wit h (0) char ge
• Almost all of mass of an at om is locat ed in t he pr ot ons and neut r ons of t he
nucleus
• The number of pr ot ons is called t he at omic number
• The t ype of at om is det er mined by t he number of pr ot ons: car bon has 6
pr ot ons, nit r ogen has 7, et c...
• Or bit s have elect r ons wit h (–) char ge
• Or bit s ar e or ganized in a ser ies of shells ar ound t he nucleus
• Elect r ons have a ver y small mass
• Chemical r eact ions involve t he elect r ons, especially t hose in t he out er shell
• Char ges in an unchar ged at om must balance (elect r ons = pr ot ons)
• I s ot op e s : if you change t he number of neut r ons in t he nucleus you pr oduce differ ent
isot opes of an element
• Some, but not all, isot opes ar e r adioact ive
• Schemat ic diagr am of 3 isot opes of car bon:
Biochemistry 63
• 6 elect r ons spin ar ound nuclei in 2 shells (inner shell has 2 elect r ons; out er has
4)
• Nuclei all have 6 pr ot ons: t his makes t hem car bon at oms
• Differ ent number s of neut r ons pr oduce differ ent isot opes:
• Car bon–12: 6 neut r ons: t his is t he most common isot ope of car bon (over
99%)
• Car bon–13: 7 neut r ons: a r ar e nat ur al isot ope
• Car bon–14: 8 neut r ons: a r adioact ive isot ope pr oduced in t he at mospher e
by cosmic r ays; it is also pr oduced in at om bomb t est s
• The isot ope number s ar e t he sums of t he pr ot ons and neut r ons and t hey
give t he r elat ive at omic masses
• Car bon–14 has a higher mass and weighs mor e t han car bon–12
• The chemical pr oper t ies of t he 3 isot opes will be almost ident ical, but some of t he
physical pr oper t ies will be differ ent
• Isot opes ar e fr equent ly used as “t r acer s” in r esear ch and medicine
VI. Molecules and Chemical bonding
• All int er act ions bet ween at oms ar e elect r ical at t r act ions bet ween char ges
• Ionic and covalent bonds hold at oms t oget her t o for m molecules
• Weak bonds (hydr ogen, van der Waals and ot her t ypes) hold molecules t oget her
• At oms make bonds by donat ing or shar ing elect r ons
• I on i c b on d s
• Compounds wit h ionic bonds split int o ions in wat er . Ions conduct elect r icit y.
Gives specialized cells (ner ve, muscle) excit able pr oper t ies.
• Suppose Na gives one of it s elect r ons t o Cl; t he Na now has a (+) char ge and t he
Cl will have a (–) char ge
• These char ged at oms ar e r efer r ed t o as ions, and since t hey have opposit e
char ges t hey at t r act each ot her and for m a chemical bond (t hey for m NaCl,
common t able salt )
• Posit ively char ged ions = cat ions (i.e, Na
+
)
• Negat ively char ged ions = anions (i.e., Cl

)
• Cova l e n t b on d s
• Each at om donat es 1 or mor e elect r ons t o t he bond
• The bonding elect r ons spend most of t heir t ime bet ween t he 2 at oms, at t r act ing
bot h nuclei and pulling t hem t oget her
• If each at om donat es 2 elect r ons t o a bond a double bond is for med
• double bonds ar e st r onger and mor e r igid t han single bonds
• A t r iple bond is for med when each at om donat es 3 elect r ons t o t he bond
• This t ype of bond holds t oget her t he long chains of macr omolecules. These
molecules do not split apar t in wat er .
64 CSIR-NET Life Sciences
• Hyd r oge n b on d s
• Occur when a hydr ogen ion is sandwiched bet ween 2 at oms, usually nit r ogen
and oxygen
• Much weaker (about 25 t imes) t han covalent or ionic bonds
• Occur bet ween molecular gr oups wit h per manent dipoles
• Wat er : makes wat er molecules st ick t oget her . Responsible for many of t he st r ange
pr oper t ies of wat er .
• Pr ot eins: cause pr ot ein chains t o spir al and bend, giving unique shapes. In DNA:
hold t oget her t he 2 chains t o for m t he double helix. Allow chains t o “unzip” for
r eplicat ion and t r anscr ipt ion.
• Va n d e r Wa a l s & ot h e r we a k b on d s
• Weak for ces t hat can bond like at oms t oget her
• Especially impor t ant bet ween chains of car bon at oms
• Alt hough weak, numer ous bonds bet ween t he chains can add up t o pr oduce
significant cohesion
• Det er mine physical st at e of compounds: gas, liquid or solid
• Occur when one at om induces a t empor ar y dipole in anot her at om
• Impor t ant in holding like molecules t oget her . Oft en det er mine t he solid, liquid
or gas st at e of a compound. Sat ur at ed fat s ar e solid at r oom t emper at ur e because
t he have mor e van der Waals at t r act ions t han unsat ur at ed fat s, which ar e liquid.
Rates of and Equilibrium States of Chemical Reactions are Determined by
Concentrations of the Reactants
• Consider a simple r eact ion in which 2 subst ances, A & B, r eact t o for m a t hir d subst ance, C:
• A + B « C
• Th e r a t e a t wh ich C is for med (in moles/secon d) will be pr opor t ion a l t o t h e
concent r at ions of A, B and C
• Rat e = Kf{A}{B} – Kb{C}
• {A}, {B} and {C} = concent r at ions of A, B and C, r espect ively; Kf = r at e const ant of
for war d r eact ion, Kb = r at e const ant of backwar d r eact ion
• For war d r eact ion r at e = Kf{A}{B}
• Backwar d r eact ion r at e = Kb{C}
• This r elat ionship is a fundament al law of chemist r y, called t he Law of Mass
Act ion
• Not e t hat if you double t he concent r at ion of eit her A or B, t he r at e of t he for war d
r eact ion will double
• Concent r at ions ar e t he number of molecules or moles per unit of volume
• Usually concent r at ions ar e given in moles/lit r e
• Exa mple: if you ha ve 3 moles of Na Cl dissolved in 6 lit er s of wa t er t he
concent r at ion is:
• 3moles/6 lit r es = 0.5 moles/lit r e = 500 mM/lit r e
• Why ar e chemical r eact ion r at es pr opor t ional t o concent r at ion?
• Molecules must collide t o r eact & number of collisions per second is pr opor t ional
t o t he concent r at ion
Biochemistry 65
• Ot her fact or s affect r eact ion r at es by changing t he r at e const ant s:
• Temper at ur e: affect s r at es in 2 ways:
• High t emper at ur e causes molecules t o move fast er pr oducing mor e collisions
• At high t emper at ur e molecules have mor e ener gy and ar e mor e likely t o
r ea ct wit h ot her molecules
• Cat alysis: r eact ion r at es can be gr eat ly acceler at ed by cer t ain t ypes of
molecules- t his will be discussed in lect ur e 4 when we consider enzymes
• A chemical r eact ion pr oceeds unt il for war d and backwar d r eact ions ar e equal
• At t his point t he r eact ion is at equilibr ium and concent r at ions do not change
• At equilibr ium:
• Kf{A}{B} = Kb{C}
• Ke = Kf/Kb = {A}{B}/{C}
• The equilibr ium const ant (Ke) is t he for war d r at e const ant (Kf) divided by
t he backwar d r at e const ant (Kb)
§ Note: at equilibrium {C} = {B}/{C}/Ke; thus, both the rates of reaction and
equilibrium concentrations are determined by concentrations of reactants
BIOENERGETICS
I. Energy producing and energy utilizing systems
A. Ca t a b oli c p a t h wa ys – gener at ion of ener gy by t r ansfor mat ion (glycolysis) or oxidat ion
(oxidat ive phosphor ylat ion) of ingest ed or st or ed fuels.
B. An a b ol i c p a t h wa ys – ut ilizat ion of ener gy for biosynt het ic pur poses (DNA synt hesis,
pr ot ein synt hesis, et c.)
C. ATP – t he link bet ween t he t wo pat hways
1. Phosphoanhydr ide bonds as high ener gy bonds
2. Ot her nucleot ides
(a) GTP: gluconeogenesis and pr ot ein synt hesis
(b) CTP: lipid synt hesis
(c) UTP: glycogen synt hesis
(d) All depend on ATP for mat ion: nucleoside diphosphat e kinase
3. Ot her bonds having a high fr ee ener gy of hydr olysis (> 7 kcal/mole)
(a) 1,3 bis-phosphoglycer at e (mixed acid anhydr ides)
(b) Phosphoenolpyr uvat e (enol phosphat es)
(c) Cr eat ine phosphat e (phosphoguanidines)
(d) Pyr ophosphat e (phosphor ic acid anhydr ides)
(e) Acet yl CoA (t hiol est er s)
II. Thermodynamic relationships and energy rich compounds
A. Laws of thermodynamics
1. F i r s t La w: ener gy may be conver t ed fr om one for m t o anot her , but t he t ot al in
a syst em r emains const ant . (Glucose ® lact at e + ATP)
66 CSIR-NET Life Sciences
2. Se con d La w: Ent r opy (DS) is a measur e of disor der or r andomness of a syst em.
All syst ems t end t o pr ogr ess t owar ds maximum ent r opy. Ent r opy is unavailable
t o per for m useful wor k.
3. Fr ee en er gy (DG): available for useful wor k
(a) DG = DH – TDS (DH =ent halpy, T = t emper at ur e in degr ees K)
(b) Exe r gon i c:fr ee ener gy lost , spont aneous r eact ion (DG less t han 0)
(c) En d e r gon i c:r equir es ener gy for r eact ion t o pr oceed (DG gr eat er t han 0)
(d) At DG = 0, t h e r e a ct i on i s a t e q u i l i b r i u m
(e) The act ual fr ee ener gy change (DG for a r eact ion is equal t o t he st andar d
fr ee en er gy (DGo’) plus a t er m t ha depends on act ual concent r at ions of
pr oduct s and r eact ant s
∆G’ =∆Go’+ RTln{[Products]/[Reactants]}
∆G’= ∆Go + 1.4 logP/ (in kcal/mole)
Where ∆Go’ = –Rtln(Keq = –1.4 log(Keq) (inkcal/mole)
(f ) F r e e e n e r gy ch a n ge s for cou p le d r e a ct i on s a r e ADDI TI VE
∆Go’ A + B ® C + D –5.0
D ® P + Q + 4.5
Sum of coupled rx A + B ® C + P + Q – 0.5
B. Thermodynamics of membrane transport
1. Unchar ged molecules: DGo’ = RT ln {[C2]/[C1]}= 1.4 log{[C2]/[C1]}
wher e t he dir ect ion of t r anspor t is fr om C1 t o C2
2. Char ged molecules: must also consider t he membr ane pot ent ial (inside -ve gener ally)
DGo’ = RT ln {[C2]/[C1]} + Z(Dy) = 1.4 log {[C2]/[C1]} + 23Z(Dy)
where F = Faraday constant = 23 kcal V–1mole–1
Dy = the membrane potential (in volts)
and Z = the charge on the ion
ENZYMES
1. Enzymes wer e discover ed by Bu ch n er (1897-1903) but t er m enzyme was given by
Ku h n e (1878).
2. Almost all enzymes ar e pr ot eins. However , r i b ozyme , r i b on u cle a s e P a r e non
pr ot ein enzyme.
3. Ever y cell pr oduces it s own enzymes because t hey cannot move fr om cell t o cell due
t o high molecular weight .
4. All component s of cell including cell wall and cell membr ane have enzymes.
5. Maximum enzymes in t he cell ar e found in mit ochondr ion.
6. Sma l l e s t e n zyme is p e r oxi d a s e and la r ge s t e n zyme being c a t a l a s e found in
per oxisomes.
7. Enzyme u r e a s e isolat ed fr om J ack bean Canavalia was cr yst allized by Summer in
1926, who pr oved pr ot ein nat ur e of enzymes.
Biochemistry 67
8. Enzymes show r ever sible r eact ions and act by lower ing ener gy of act ivat ion by mor e
t han 50%.
9. Enzymes show t hr ee dimensional st r uct ur es.
10. K
m
(Mi ch a e l i s Me n t i on Con s t a n t ) is t he subst r at e concent r at ion at which t he
chemical r eact ion at t ains half it s maximum velocit y. It is an inver se measur e of t he
affinit y of an enzyme for it s subst r at e. Small t he K
m
, t he gr eat er t he subst r at e affinit y.
Allost er ic enzymes do not obey K
m
const ant .
11. Over 2000 enzymes have been r ecor ded. Enzymes ar e synt hesized by living cells.
Most of t h e en zymes r ema in a n d fu n ct ion in side t h e cells. Th ese a r e ca lled
en d oen zymes (or i n t r a ce llu la r e n zyme s ). On t he ot her hand, t he enzymes which
leave t he cells and funct ion out side t hem ar e called exoen zymes (or e xt r a ce l l u l a r
en zymes ). These r et ain t heir cat alyt ic abilit y even when ext r act ed fr om cells. Rennet
t ablet s (cont aining t he enzyme r ennin fr om t he calf’s st omach) have been in use for
coagulat ing milk pr ot ein t o obt ain casein (cheese fr om milk).
12. A lar ge number of enzymes r equir e an addit ional non pr ot ein component called
pr ost het ic gr oup for t heir efficient act ivit y. Pr ost het ic gr oup may be divided r at her
loosely int o t wo gr oups – (a) Me t a l a ct i va t or s ; (b) Cofa ct or s or coen zymes .
13. In h oloe n zyme s , t he a p oe n zyme (pr ot ein par t ) det er mines t he specificit y while
coe n zyme (pr ost het ic gr oup) det er mines t he cat alyt ic funct ional act ivit y of enzyme.
14. The pr ot einaceous par t of enzyme is called a p oe n zyme. The apoenzyme plus non
pr ot einaceous par t is called h ol oe n zyme. Some enzymes r equir e a loose associat ion
wit h cer t ain or ganic subst ances for t heir act ivit y. These pr ost het ic gr oups ar e called
cofa ct or s or coen zymes . Examples of coenzymes ar e – NAD (nicot inamide adenine
dinucleot ide), NADP (nicot inamide adenine dinucleot ide phosphat e), ATP (adenosine
t r iphosphat e), CoA (coenzyme A), F MN (Flavin mononucleot ide) and FAD (Flavin
adenine dinucleot ide). FAD and FMN cont ain r i b ofla vi n (vit amin B
2
) as a component .
Ri b ofl a vi n is t he hydr ogen accept ing par t of FAD/FMN.
15. Enzymes ar e r equir ed in minut e quant it ies which is sufficient t o conver t a lar ge
amount of subst r at es (st ar t ing mat er ials of a r eact ion) t o pr oduct s (ending mat er ials
of a r eact ion).
16. Ever y enzyme has it s own opt imum pH. Any shift t owar ds alkaline or acidic side
r esult s in a decr ease in enzyme act ivit y because it denat ur es t he enzyme molecule
(changes it s shape). P e p s i n of gast r ic juice has opt imum act ivit y at pH 2.0, while
t r yp s i n shows maximum acit ivit y at pH 8.0.
17. Ever y enzyme has a specific opt imum t emper at ur e.
18. Over a r ange of 0–40°C, t he r at e of enzyme cont r olled r eact ion almost doubles for
ever y r ise of 10°C. (Q
10
= 2).
19. Most enzymes show maximum act ivit y in a t emper at ur e r ange of 25–40°C.
20. Enzymes ar e t her molabile i.e. ar e denat ur ed at high t emper at ur es. The loss of cat alyt ic
pr oper t ies begins at 36°C and is almost complet e as 60°C is r eached. However , dr ied
enzyme ext r act s can endur e t emper at ur e of 100°C–120°C or even higher . That is
why; dr y seeds can endur e high t emper at ur e t han ger minat ing seeds.
21. Enzymes ar e gener ally specific for t he t ype of r eact ions t hey cat alyse. This specificit y
is ver y st r ong for some enzymes.
68 CSIR-NET Life Sciences
22. Enzymes ar e colloidal in nat ur e; and have high molecular weight s r aning fr om 10,000–
50,000. However , t he molecular weight s of cat alase and ur ease ar e 2,50,000 and
4,83,000 r espect ively.
23 I UB (1962) has divided enzymes int o 6 classes (oxi d or e d u ct a s e s , t r a n s fe r a s e s ,
h yd r ol a s e s , l ya s e s , i s ome r a s e s and li ga ses ). Each class is divided int o sub-classes
and each sub-class int o sub-sub classes depending upon t he t ype of r eact ion and nat ur e
of subst r at e. Thus ever y enzyme has a four digit code called EC Number (Enzyme
Commission Number ).
24. Th e e n zyme s a r e cl a s s i fi e d i n t o 6 gr ou p s on t h e b a s i s of t yp e of r e a ct i on s
t h e y ca t a l ys e:
(a) Oxi d or e d u ct a s e s : Tr ansfer of H and O at oms or elect r ons fr om one subst ance
t o anot her . Examples ar e dehydr ogenase, oxidase.
(b) Tr a n sfe r a se s : Tr ansfer of a specific gr oup (met hyl, aceyl, amino or phosphat e)
fr om one subst ance t o anot her . Examples: t r ansaminase, kinase.
(c) Hyd r ol a s e s : Hydr olysis of a subst r at e. Examples ar e lipase, amylase, pept idase,
est er ase, phosphat ase, car bohydr ase, and pr ot ease.
(d) Lya ses : Non hydr olyt ic r emoval or addit ion of gr oup fr om subst r at es, C–C, C–
N, C–O, or C–S bonds may be split . Examples ar e decar boxylase, fumar ase, and
aldolase.
(e) I s ome r a s e s : Cha nge of a subst r a t e int o a r ela t ed for m by int r a molecula r
r ear r angement . Examples ar e phosphohexose isomer ase.
(f) Li ga s e s (s yn t h e t a s e s ): J oining of t wo molecules by synt hesis of new C–O, C–
S, C–N or C–C bonds wit h simult aneous br eakdown of ATP. Examples ar e acet yl
CoA synt het ase (act ing on fat t y acids), pyr uvat e car boxylase.
25. The enzyme lower s t he act ivat ion ener gy of a r eact ion. (The ener gy r equir ed for
subst r at es t o r eact in or der t o get conver t ed int o pr oduct is called ener gy of act ivat ion).
An enzyme (E) combines wit h it s subst r at es (S) t o for m a shor t -lived enzyme-subst r at e
(ES) complex. Wit hin t his complex, t he chances of occur r ing of r eact ion ar e gr eat ly
incr eased. Once a r eact ion has occur r ed, t he complex br eaks up int o pr oduct s and
enzymes. Thus, t he enzyme r emains unchanged at t he end of t he r eact ion; and is
fr ee t o int er act again wit h mor e subst r at es.
26. Most enzymes ar e far lar ger molecules t han subst r at es. Only a small por t ion of t he
enzyme (3-12 amino acids) comes int o dir ect cont act wit h t he subst r at e. This r egion
is called t he act ive sit e of t he enzyme. An enzyme may have mor e t han one act ive
sit e. The r emaining amino acids maint ain t he cor r ect globular shape of t he molecule.
It is impor t ant for pr oper funct ioning of t he a c t i ve s i t e.
Fisch er (1890) proposed lock a n d k e y h yp ot h e s i s t o explain specificit y. He pr oposed
t hat enzymes have a par t icular shape int o which t he subst r at e or subst r at es fit exact ly.
For a lock t o wor k, it must be pr ovided wit h r ight key. Similar is t he case wit h
enzyme and subst r at es.
27. The best evidence for t his lock and key hypot hesis (or t emplat e t heor y) of enzyme
act ion comes fr om t he obser vat ion t hat compounds similar in st r uct ur e t o t he subst r at e
inhibit t he r eact ion.
28. Evidence fr om pr ot ein chemist r y suggest ed t hat a slight r ear r angement of chemical
gr oups occur s in bot h enzyme and subst r at e when as ES complex is for med. It means
Biochemistry 69
t hat enzymes and t heir act ive sit es ar e r at her flexible st r uct ur es. Kos h l a n d (1959),
t her efor e, suggest ed induced-fit hypot hesis. Accor ding t o it , when a subst r at e combines
wit h a n enzyme, it induces cha nge in t he enzyme st r uct ur e. The a mino a cids
const it ut ing t he act ive sit e ar e moulded int o a pr ecise for mat ion which enables t he
enzyme t o per for m it s cat alyt ic funct ion mor e effect ively.
29. En zyme a ct i on ca n b e i n h i b i t e d b y (a) denat ur at ion of enzymes (b) compet it ive
i n h i bi t i on (c) non compet it ive inhibit ion (d ) a llost er ic modifica t ion of feedba ck
inhibit ion.
(a) De n a t u r a t i on : Change in t he spat ial ar r angement of polypept ide chain wit hin
t he pr ot ein molecule so t hat it s unique st r uct ur e is changed. As a r esult , t he
physical or biological pr oper t ies ar e changed.
(b) Re ve r s i b le I n h i b i t i on :
1. Comp e t i t i ve i n h i b i t i on : A subst a nce which closely r esembles a ct ua l
subst r at e st r uct ur e compet es wit h t he subst r at e for t he act ive sit e. Ther e
is a decline in t he number of act ual subst r at e molecule binding t he sit e, as
many of t hese ar e occupied by closely r esembling subst ance. As a r esult ,
enzyme act ion is inhibit ed. Can be r ever sed by incr easing t he subst r at e
concent r at ion. Vma x u n a ffe ct e d & Ap p a r e n t Km i n cr e a s e d by fact or
(1 + [I]) /KI). Exa mp l e s :
• I nhibit ion of a ct ivit y of succina t e dehydr ogena se by ma lona t e &
oxaloacet at e.
• Sulpha dr ugs e.g. sulphanilamide inhibit t he synt hesis of folic acid in
bact er ia by compet ing wit h p-amino benzoic acid (PABA).
• AZT and AIDS
• Met hot r exat e and dihydr ofolat e r educt ase
2. Non comp e t i t i ve i n h i b i t i on : The inhibit or has no st r uct ur al similar it y
t o t he subst r at e and for ms an enzyme-inhibit or complex at a point ot her
t han it s act ive sit e so t hat t he globular st r uct ur e of t he enzyme is changed.
As a r esult , even if t he subst r at e is able t o bind wit h t he enzyme, cat alysis
cannot t ake place. It is char act er ist ic of allost er ic and mult isubst r at e
enzymes. Km u n a ffe ct e d in simplest case. Vma x d e cr e a se d by fact or (1
+ [I])/KI) Cyanide inhibit s t he act ivit y of cyt ochr ome oxidase. Di isopr opyl
fluor ophosphat es (ner ve gas) effect s ner ve impulse t r ansfer by combining
ir r ever sibly wit h amino acid ser ine of acet ylcholine est er ase. It also poisons
a number of ot her enzymes wit h t r ypsin, chymot r ypsin, phsphoglucomut ase,
elast ase et c. Iodo acet amide inhibit s enzymes having sulphahydr ol or
imidazole gr oup.
3. Un comp e t i t i ve (I binds t o ES complex). Km, Vma x b ot h d e cr e a s e.
Char act er ist ic of mult isubst r at e enzymes
(c) I r r e ve r s i b l e i n h i b i t or s : Gener ally achieved by Covalent modificat ion. Similar
t o non-compet it ive kinet ics (Vma x l owe r e d , Km u n a ffe ct e d ) Examples:
(i) PCMB and GAPDH
(ii) DFP and Ser ine hydr olases (t r ypsin, chymot r ypsin, acet yl choline est er ase,
et c)
(iii) Suicide subst r at es: gener at ion of r eact ive int er mediat e by enzyme
70 CSIR-NET Life Sciences
(d) F e e d b a ck i n h i b i t i on : The pr oduct of an enzyme cat alysed r eact ion/chain of
r eact ions accumulat es and act s as inhibit or of t he r eact ion. e.g. Inhibit ion of
t hr eonine dea mina se by isoleucine. Pr ot ea ses a r e a dded in det er gent s a nd
amylase is used for dish washing.
• Tr yp s i n is added t o par t ially pr edigest ed food. Immobilisat ion of enzymes
is done by at t aching or t r apping enzymes in iner t suppor t ing mat er ials for
bet t er efficiency and r ecover ing t hem aft er t he r eact ion.
• Alloe n zyme s ar e enzymes which ar e pr oduced by differ ent genes.
• Con s t i t u t i ve e n zyme s a r e a lwa ys pr esent beca use t hey a r e a lwa ys
r equir ed for vit al pr ocess e.g. glycolysis.
• Re p r e ssi b le e n zyme s – nor mally r emain pr esent but ar e r epr essed when
a specific chemical or pr oduct is pr esent e.g. glucokinase.
• I n d u ci b le en zymes – ar e for med in r esponse t o pr esence of it s subst r at e
e.g. la ct ose.
• ELI SA – It is an enzyme linked immunosor bent assay when a pr ot ein,
ant ibody or ant igen is det ect ed by means of a specific enzyme e.g. AIDS.
• Re s t i ct i on e n d on u cl e a s e – These ar e enzymes which ar e used t o br eak
DNA a t a specific sit e pr oducing st icky ends. The enzymes a r e highly
impor t ant for genet ic engineer ing. Ar ber , Nat hans and Smit h wer e awar ded
Nobel Pr ize in 1978 for t heir discover y.
ENZYME KINETICS
A. Qu a n t i t a t i on of e n zyme s
1. a ct i vi t y = “h ow mu ch ” ( moles pr oduct for med per minut e)
2. s p e ci fi c a ct i vi t y = “h ow p u r e ” ( moles pr oduct /min. per mg pr ot ein)
3. Tu r n ove r n u mb e r , k ca t = “h ow e ffi ci e n t ” = Vma x/[ET] (moles pr oduct /min.
per mole enzyme)
B. I n i t i a l ve l oci t y p l ot s
1. v
o
vs [Enzyme] assumed linear
2. v
o
vs [Subst r at e] non-linear
C. As s u mp t i on s for d e r i va t i on of Mi ch a e l i s Me n t e n e q u a t i on
1. Mechanism E + S <——> ES —> E + P
**2. [ES] const ant (“st eady st at e”)
3. [S] >> [Enzyme]
4. [P] ~ O (init ial velocit y) Not r esponsible for der ivat ion of Michaelis-Ment en
equat ion
D. Me a n i n g of Km, Vma x, k ca t
1. Km is t he concent r at ion of subst r at e r equir ed t o achieve 1/2 Vmax.
2. kcat , x [Enz] = Vmax
E. De t e r mi n a t i on of Km, Vma x Lineweaver -Bur k plot s: 1/vo vs 1/[S]
Biochemistry 71
V. Multi-substrate enzyme mechanisms
A. Sequent ial: all subst r at es combine befor e any pr oduct r elease
B. Ping pong: pr oduct r elease befor e all subst r at es combine
ENZYMES: CONTROL OF ENZYME ACTIVITY
I. Overview of factors influencing enzyme activity
A. Envir onment al fact or s
1. Temper a t ur e
2. Ionic st r engt h
3. pH
4. Concent r at ions of subst r at es (r eact ant s), pr oduct s, cofact or s
B. Rat e of synt hesis and degr adat ion of enzyme (minut es t o hour s)
C. I s ozyme s : mult iple for ms of an enzyme cat alyzing t he same r eact ion in t he
same or ganism
D. Covalent modificat ion of enzyme (seconds t o minut es)
E. Allost er ic r egulat ion (seconds)
F. Pr ot eolyt ic cleavage:
1. Act ivat ion: zymogens ar e inact ive enzyme pr ecur sor s t hat ar e act ivat ed by
pr ot eolyt ic cleavage (e.g. digest ive enzymes t r ypsin, chymot r ypsin, et c.)
2. Inact ivat ion by pr ot eolysis: also digest ive enzymes
G. Binding of a r egulat or y pr ot ein: e.g. t r ypsin and t r ypsin inhibit or
H. Compar t ment at ion:
1. Met abolic channeling: dir ect t r ansfer of subst r at e t o next enzyme (Pyr uvat e
dehydr ogenase complex)
2. Separ at e subcellular locat ion of cat abolic, anabolic pat hways (e.g. fat t y acid
oxidat ion in mit ochodr ia, synt hesis in cyt osol)
II. Isozymes: different enzymes catalyzing the same reaction
A. Hexokinase and Glucokinase
(The br ain enzyme is nor mally sat ur at ed wit h subst r at e due t o it s low Km)
B. Lact ic dehydr ogenase (LDH)
1. Five (5) isozymes possible (t et r amer wit h 2 t ypes of subunit s, H & M)
2. Isozyme pat t er n var ies wit h t issue
3. Tissue damage r esult s in leakage int o ser um
Type Composition Location
LDH1 HHHH Myocardium and RBC
LDH2 HHHM RBC and myocardium
LDH3 HHMM Brain and kidney
LDH4 HMMM
LDH5 MMMM Liver and skeletal muscle
72 CSIR-NET Life Sciences
C. Cr eat ine (phospho) kinase (CPK or CK)
1. A dimer of M and B subunit s for ms 3 separ at e isozymes (MM,MB,BB)
2. Diagnosis of myocar dial infar ct s (MI)
III. Zymogens and Proenzymes: inactive precursors of enzymes
A. Nomencla t ur e
1. Tr ypsinogen – inact ive pr ecur sor of t r ypsin
2. Pr ocar boxypept idase – inact ive pr ecur sor of car boxypept idase
B. Act ivat ion by pr ot eolysis (e.g., digest ive enzymes fr om pancr eas)
C. Role of Tr ypsin inhibit or : block pr emat ur e act ivat ion
D. Specificit y compar ison T, CT, Elast ase
IV. Regulation of enzyme activity by metabolic intermediates
A. Br anch point s in met abolism A® B « C ® D «


E ®F « G
B. Commit t ed st ep (fir st ir r ever sible in specific pat h)
C. Feedback inhibit ion/cr oss r egulat ion
D. Classic r egulat or y enzyme is ALLOSTERIC
V. Properties of allosteric enzymes
A. Unusual kinet ics
1. Classic example is “sigmoid” kinet ics (S-shaped V
o
v/s [S] cur ve) which is
r efer r ed t o as posit ive cooper at ivit y bet ween subunit s
2. Many ot her possibilit ies: at ypical r at e vs [subst r at e] cur ves.
B. Usually Mult i-subunit , Separ at e binding sit e (allost er ic sit e) for act ivat or s, inhibit or s
(posit ive or negat ive modulat or s or effect or s)
C. Example: Pr ot ein kinase A
1. cAMP binds t o r egulat or y (R) subunit s
2. Cat alyt ic subunit s r eleased, act ive
3. Cat alyt ic and Regulat or y sit es ar e on separ at e polypept ides
D. Example: aspar t at e t r anscar bamoylase
1. ATP (subst r at e) a posit ive effect or
2. CTP (end pr oduct of pat hway) a negat ive effect or
3. Cat alyt ic and Regulat or y sit es on separ at e polypept ides
VI. Models for allosteric enzymes
A. The Concer t ed (Symmet r y) model: R « T
Biochemistry 73
B. The Sequent ial or induced fit model: block of 4 post age st amps
C. Bot h involve Tense and Relaxed subunit confor mat ions
VII. Regulation by phosphorylation (covalent modification)
A. Ser ine, t hr eonine, t yr osine hydr oxyls
B. Example: Glycogen phosphor ylase
ENZYMES: MECHANISMS OF ACTION
I. Introduction and terms
Nucleophile, elect r ophile,
car bonium ions, oxonium ions, car banions,
t r ansit ion st at e, int er mediat e,
act ivat ion ener gy
II. Active site: often polar and ionizable amino acids important
A. Cat alyt ic cent er
B. Binding sit e
III. Theoretical explanations for enzyme catalysis
A. “Lock and Key”
1. Pr oximit y - “local concent r at ion”
2. Or ient at ion effect s - alignment of bonds t o facilit at e cat alysis
B. Induced fit - Hexokinase (enzyme confor ms t o subst r at e)
C. Induced st r ain - Lysozyme (subst r at e confor ms t o enzyme )
D. Tr ansit ion st at e st abilizat ion (TSS)
1. Tr ansit ion st at e analogs ar e excellent enzyme inhibit or s
2. Dr ug design based on Tr ansit ion St at e analogs
3. Ab zyme s : ant ibodies against t r ansit ion st at e analogs have enzymat ic act ivit y
IV. Enzymatic catalysis mechanisms
A. Aci d -b a s e ca t a l ys i s
1. Gener a l model Enz–H
+
+ R' – OR ® Enz + R' – H + R
+
R
+
+ H
2
O + Enz ® Enz–H + R – O –H
2. Specific examples: (a) Lysozyme (b) Ribonuclease
B. Cova l e n t ca t a l ys i s : i n t e r me d i a t e wi t h cova l e n t b on d b e t we e n E a n d S
1. Gener al model: Enz–O– H + A–B ® Enz–O–A + B– H
Enz–O–A + H
2
O ® Enz – O – H + A – O –H
2. Specific examples:
(a) Chymot r ypsin
(b) Subt ilisin: conver gent evolut ion
74 CSIR-NET Life Sciences
C. Me t a l i on ca t a l ys i s
1. Funct ion as Lewis acids
2. Funct ion t o st abilize int er mediat es (chelat es)
3. Facilit at e binding by neut r alizing char ges
4. Involved in many r edox r eact ions
LIPIDS
Li p i d s ar e est er s of fat t y acids and polyhydr ic alcohol. The t er m ‘lipid’ was fir st used by
Bl oor (1943). These ar e t he compounds of C, H, O but t he r at io of H and O is mor e t han
2:1 (i.e., t he r at io of oxygen is lesses as compar ed t o car bohydr at es). These ar e wat er -
insoluble or ganic subst ances which can be ext r act ed fr om t he cells by or ganic solvent s
such as ether, chloroform and benzene. Their general form ula is C
n
H
2n
O
2
. Some lipids
have P, N and S also.
1. Si mp l e l i p i d s – est er s of fat t y acids wit h alcohol. Simplest alcohol in fat s is glycer ol
(a t r ihydr ic alcohol) e.g., fat s, oils and waxes. Tr iglycer ides ar e common in nat ur e.
2. Comp ou n d l i p i d s – These lipids cont ain an addit ional gr oup alongwit h fat t y acids
and alcohols, e.g., phospholipids, glycolipids and lipopr ot eins.
3. De r i ve d li p i d s – These ar e isopr enoid st r uct ur es e.g., st er oids, t er penes, car ot enoids.
• F a t t y a ci d s ar e car boxylic acid wit h a chain of mor e t han four car bon at oms ending
wit h COOH gr oup. Plant s can synt hesize all fat t y acids. Animals can not synt hesize
li n ole i c, li n ole n i c and a r a ch i d on i c a ci d . These ar e called e s s e n t i a l fa t t y a ci d s .
Their deficiency causes st er ilit y, kidney failur e and st unt ed gr owt h.
• Sat ur at ed fat t y acids have no double bond. Their melt ing point is high. Palmit ic acid,
st ear ic acid ar e sat ur at ed fat t y acids.
• Unsat ur at ed fat t y acids ar e commonly pr esent in veget able oils, cod/shar k oil. Their
melt ing point s ar e low. Oleic acid has one double bond, linoleic acid has t wo, linolenic
acid has t hr ee and ar achiodonic acid has 4 double bonds. Fat t y acids wit h mor e t han
one double bond ar e called p ol yu n s a t u r a t e d fa t t y a ci d (P UF A).
• Dr yi n g oi l s ar e unsat ur at ed fat t y acids which can be conver t ed in har d fat s on
being exposed.
• Edible oils can be conver t ed int o har d fat s t hr ough h yd r oge n a t i on .
• Waxes ar e est er s of long chain monohydr ic alcohols like ce t yl, ce r yl or me r i cyl .
• La n oli n for ms a pr ot ect ive, wat er insoluble coat ing on animal fur .
• Bees wa x secr et ed fr om abdominal glands of honey bees has p a l mi t i c a ci d
and me r i cyl a lcoh ol.
• P a r a ffi n wa x is a pet r oleum pr oduct .
• Cu t i n is for med by cr oss est er ificat ion and polymer izat ion of hydr oxyl fat t y
acids and ot her fat t y acids wit hout est er ificat ion by alcohols ot her t han glycer ol.
Cut icle has 50-90% cut in.
• Su ber i n is condensat ion pr oduct of gl yce r ol and p h e l l on i c a ci d . It makes t he
cell wall imper meable t o wat er .
Biochemistry 75
• P h os p h ol i p i d s ar e t r iglycer ides in which one fat t y acid is r eplaced by phosphor ic
acid which is oft en linked t o addit ional nit r ogenous gr oups like ch ol i n e (in lecit hin),
e t h a n ol a mi n e (in cepalin), s e r i n e or i n os i t ol . These ar e amphipat hic i.e. have
bot h polar and non polar gr oups. These for m cell membr anes along wit h pr ot eins.
• Sp h i n gol i p i d s have amino alcohol sphingosine. Sp h i n gomye l i n s ar e pr esent in
myelin sheat h of ner ves. They have addit ional phosphat e at t ached t o choline, ar e
pr esent in ner ve membr ane. Cer ebr osi d es ar e pr esent in ner ve membr ane and
have ga l a c t os e.
• Ga n gl i os i d e s – ha ve glucose, ga la ct ose, s i a l i c a ci d and a ce t yl gl u cos a mi n e,
pr esent in gr ey mat t er , r ecept or s of vir al par t icles, excess causes Ta y-Sa ch s disease.
• St e r ols or s t e r oi d s cont ain 4 fused hydr ocar bon r ings called cyclop en t a n e p er h yd r o
p h e n a n t h r e n e and a long side chain e.g. cholest er ol, st igmast er ol, campest er ol,
sit ost er ol, er gast er ol.
• Cholest er ol helps in absor pt ion of fat t y acids, sex hor mones, vit amin D and bile
salt s. Pot at o is r ich in cholest er ol. Excess of cholest er ol causes a t h e r osch e r osi s.
• P r os t a gl a d i n s ar e hor mone modulat or s.
• Ter penes ar e lipid like car bohydr at es for med of isopr ene unit s e.g., ment hol, camphor ,
car ot enoids.
• Major funct ion of lipids is t o act as ener gy st or es.
• Plant s usually st or e oil t han fat s. Seeds, fr uit s and cholor oplast s ar e oft en r ich
in oils.
• Gl y c o l i p i d s a r e i mpor t a n t compon en t s of cel l membr a n es , ch l or opl a s t
membr a nes.
• Fat t y subst ance in t he cell wall (Wax, cut in, suber in) helps r educe t r anspir at ion
and pr ovide mechanical pr ot ect ion fr om injur y and par asit es.
• Di os ge n i n is a st er oid obt ained fr om t he plant called Dioscorea. It is used for
manufact ur ing ant ifer t ilit y pills.
MEMBRANE STRUCTURE AND TRANSPORT
I. Functions of membranes
Per meabilit y bar r ier , t r anspor t , communicat ion wit h t he “out side”, cell-cell r ecognit ion,
envir onment for non-aqueous r eact ions, cell volume r egulat ion and locomot ion.
II. Chemical composition of lipids
A. Fat t y acid st r uct ur es:
1. Be able t o ident ify st r uct ur es of common fat t y acids p a l mi t i c, s t e a r i c,
ol e i c, l i n ol e i c, l i n ol e n i c a n d a r a ch i d on i c.
2. Double bonds and shor t er chains lower t he melt ing point
B. Membr ane lipids: amphipat hic st r uct ur es
1. Gl yce r op h os p h ol i p i d s
(a) Phosphat idat e + “X” (PC, PE, PS, PG, PI, and car diolipin-)
76 CSIR-NET Life Sciences
(b) Plasmalogens (Glycer ol et her phospholipids)
(i) Plat elet act ivat ing fact or (PAF): acet yl at R2 and choline at R3
(ii) Lower s blood pr essur e, causes plat elet s t o aggr egat e
(c) Lysophosphoglycer ides have R2 vacant ; det er gent act ion
2. Sp h i n gol i p i d s
(a) Sphingosine as par ent al compound
(b) Ce r a mi d e s : N-fat t y acyl sphingosine
(c) Ce r e b r os i d e s : cer amide + glu or gal
(d) Su l fa t i d e s : gal-cer ebr oside t hat is sulfat ed
(e) Ga n gl i os i d e s : cer amide + br anched sugar , including N-Acet yl-Neur aminic
Acid (NANA or sialic acid).
(i) Ganglioside GM1 is t he r ecept or for choler a t oxin
(ii) Var iet y of Sphingolipid st or age diseases (Tay Sachs, et c.)
3. Sp h i n g o my e l i n : a s ph in go-ph os ph o-lipid (Not e s t r u ct u r a l s imila r it y t o
glycer ophospholipids)
4. Ch ol e s t e r ol : impor t ant component of membr ane; makes membr ane less fluid
& pr ecur sor t o st er oid hor mones
III. Properties of lipid aggregates
A. Mi ce lle s : spher ical aggr egat es of single-t ailed lipid
B. Li p i d b i l a ye r s : aggr egat es of double-t ailed phospholipids
1. Li p os ome s : pr oposed use as dr ug deliver y agent s
pr oper t ies:
(i) imper meant t o wat er solut es
(ii) lat er al diffusion r apid
(iii) t r ansver se (flip-flop) mot ion RARE
(iv) fluidit y decr eases wit h chain lengt h, sat ur at ion of FA
2. Planar Bilayer s: similar pr oper t ies
IV. Biological membranes
A. The Fluid Mosaic Model of Membr anes by Sanger and Nicolson (t wo dimensional
asymmet r ic lipid bilayer wit h pr ot eins suspended wit hin, car bohydr at e only at t ached
on out side of membr ane).
B. Me mb r a n e p r ot e i n s :
1. P e r i p h e r a l or e xt r i n s i c
(a) asymmet r ic in dist r ibut ion
(b) Var iet y of membr ane anchor s
(i) ionic int er act ions (cyt ochr ome c of mit ochondr ia)
(ii) hydr ophobic amino acid sequence (cyt o. b5 of E.R.)
(iii) GPI anchor s (glycosyl phosphat idylinosit ol)
Biochemistry 77
(iv) Myr ist ic (C–14) via amide t o C-t er minal glycine
(v) Palmit ic (C–16) acid via t hioest er t o cys
(vi) Pr enylat ed or isopr enoid gr oups: far nesyl (C–15) or ger anyl- ger anyl
(C–20) at C-t er minal cys
2. I n t e gr a l or i n t r i n s i c
(a) t r ansmembr ane helices (>20 hydr ophobic amino acids)
(b) specific or ient at ion (N, C t er mini)
(c) examples: glycophor in (one t r ansmembr ane helix); anion channel of RBC
(12 transmembrane helices) (Cl

/HCO
3

excha nger )
(d) Por ins – Tr ansmembr ane - pleat ed sheet mit ochondr ia, bact er ia)
RARE: almost always alpha-helix acr oss membr ane
C. Ca r b oh yd r a t e – ext er nal side of plasma membr ane
1. At t ached t o pr ot ein side chains
(a) O-l i n k e d : SER, THR (wit hin polypept ide)
(b) N-l i n k e d : ASN (wit hin polypept ide)
2. Car bohydr at e is impor t ant for “t ar get ing” pr ot eins t o subcellular
compar t ment s
D. Cholest er ol wit h –OH at sur face, r ings wit hin membr ane (decr eases fluidit y)
V. Properties of biological membranes
A. Me mb r a n e flu i d i t y
1. Lipids: lat er al mot ion r apid, t r ansver se r ar e; asymmet r ic locat ion
2. Pr ot eins: lat er al mot ion var iable (cyt oskelet on int er act ions), t r ansver se r ar e
3. Gener al bar r ier t o wat er -soluble molecules, but not t o wat er
B. Er yt hr ocyt e (RBC) membr ane as example
1. RBC a “dying cell”; no nucleus, no mit ochondr ia (120 days lifet ime)
2. Membr ane r olls fr eely ar ound cell yet cell maint ains discoid shape
3. Membr ane pr ot eins
(a) Glycophor in A
(b) “Band 3” is HCO
3

/Cl

anion exchange pr ot ein
(c) Band 3 at t ached t o Ankyr in, which in t ur n is at t ached t o Spect r in, which
ar e bot h par t of cyt oskelet on
(d) Cyt oskelet on limit s mot ion of cer t ain at t ached membr ane pr ot eins
(e) Her edit ar y ellipt ocyt osis – defect ive spect r in - malar ial belt
VI. Membrane transport of small molecules (nutrients), ions
A. Ch a r a ct e r i s t i cs of me mb r a n e t r a n s p or t : pr ot ein mediat ed, specific, inhibit able,
exhibit sat ur at ion kinet ics like enzymes.
B. Typ e s of me mb r a n e t r a n s p or t
1. Si mp l e P a s s i ve Di ffu s i on
(a) not sat ur able, not specific, not inhibit able
(b) depends on solubilit y in membr ane (non-polar pr efer r ed)
78 CSIR-NET Life Sciences
2. F a ci l i t a t e d d i ffu s i on : Pr ot ein mediat ed (t r anspor t er s, t r anslocases, por t er s,
car r ier s, per meases)
(a) sat ur at ion kinet ics, specific, specific inhibit or s
(b) ACTI VE TRANSP ORT – movement AGAINST t he elect r ochemical gr adient
(i) pr imar y – uses ATP as ener gy
(ii) secondar y – uses ion gr adient s for ener gy (which in t ur n r equir e ATP)
(c) P ASSI VE TRANSP ORT – movement DOWN t he concent r at ion gr adient )
(i) channels (specific) or por es (non-select ive) – membr ane hole, no binding
of molecule; somet imes ga t ed
(ii) mediat ed t r anspor t – specific binding by pr ot ein involved
(d) mode of t r ansmembr ane movement
(i) Unipor t – molecule moves by it self
(ii) Sympor t – molecule moves along wit h anot her molecule
(iii) Ant ipor t – molecule moves opposit e dir ect ion fr om anot her
(iv) Elect r oneut r al – no net change in t r ansmembr ane char ge
(v) Elect r ogenic – t r anspor t alt er s t r ansmembr ane char ge
C. Th e r mod yn a mi cs of me mb r a n e t r a n s p or t
1. An elect r ical pot ent ial acr oss membr anes (“membr ane pot ent ial”) ar ises fr om
differ ences in per mea bilit y a nd concent r a t ion for va r ious ions a cr oss t he
membr a ne.
2. The ener gy r equir ed t o move a molecule acr oss a membr ane depends on:
(a) t he t r ansmembr ane concent r at ion gr adient of t hat molecule, and
(b) t he membr ane pot ent ial if t he t r anspor t pr ocess is Elect r ogenic
+ Z(d e l t a P s i )F (i n k ca l /mol e )
wher e F = 23 kcal V–1 mol–1 (Far aday const ant ),
Z is t he net char ge moved, and (delt aPsi) is t he membr ane pot ent ial in
volt s.
D. Sp e ci fi c e xa mp l e s of t r a n s p or t s ys t e ms
1. P a s s i ve me d i a t e d : glucose t r anspor t int o muscle, adipose t issue
(a) glucose binds on one sur face
(b) confor mat ional change occur s, exposing glucose t o t he opposit e side of t he
membr ane wher e it is r eleased
(c) no ener gy input , moves DOWN conc. gr adient
(d) Insulin st imulat es movement of t r anspor t er pr ot ein fr om int er nal vesicle
t o membr a ne
2. Act i ve t r a n s p or t
(a) Pr ima r y: Na
+
K
+
ATPase of plasma membr ane
(i) ATP hydr olysis is dir ect ly involved
(ii) bot h Na
+
and K
+
move against t he elect r ochemical gr adient
(iii) Phosphor ylat ion of specific Asp t r igger s confor mat ional change causing
INTERNALLY bound Na
+
t o be r eleased OUTSIDE cell
(iv) Dephosphor yla t ion of Asp ca uses EXTERNALLY bound K
+
t o be
r eleased INSIDE cell
Biochemistry 79
(b) Secondary: N a
+
glucose sympor t int o int est inal epit helium
(i) ATP only INDIRECTLY involved (t o maint ain Na
+
gr adient )
(ii) Na
+
gr adient supplies t he ener gy for glucose ent r y
(iii) Glucose exit s epit helial cell int o t he cir culat or y syst em via passive mediat ed
sympor t
(iv) Amino acids t r anspor t ed similar ly
E. I on op h or e s : bact er ial poisons of ion t r anspor t syst ems
1. Mobile car r ier s: Valinomycin for ms cage for K
+
2. Channel for mer s: Gr amicidin for small inor ganic ions; r elat ively non-specific
CARBOHYDRATES
Ca r b oh yd r a t e s ar e mainly compounds of car bon, hydr ogen and oxygen. Car bohydr at es ar e so
called because in most of t hem, t he pr opor t ion of hydr ogen and oxygen is t he same as in wat er
(H
2
O) i.e., 2:1. Their gener a l for mula is C
X
H
2X
O
X
. These a r e a lso knows a s sa ccha r ides
(compou n ds con t a i n i n g s u ga r ). Ca r boh ydr a t es a r e pr odu ced by gr een pl a n t s du r i n g
phot osynt hesis. These const it ut e about 80% of t he dr y weight of plant s. Car bohydr at es ar e
di vi ded i n t o 3 ma i n cl a s s es – m o n o s a c c h a r i d e s , d e r i v e d m o n o s a c c h a r i d e s a n d
oli gos a cch a r i d e s .
1. Monosaccharides
• These have single sacchar ide unit s which cannot be hydr olysed fur t her int o st ill
smaller car bohydr at es; have gener al for mula C
N
H
2N
O
N.
These ar e composed of 3–7
car bon at oms, and ar e classified accor ding t o t he number of C at oms as t r ioses (3C),
t et r oses (4C), pent oses (5C), hexoses (6C) and hept oses (7C). Of t hese, pent oses and
hexoses ar e most common. Monosacchar ides ar e impor t ant as ener gy sour ce and as
building blocks for t he synt hesis of lar ge molecules.
• All monosacchar ides ar e eit her aldoses or ket oses. The t wo simplest monosacchar ides
ar e t r ioses e.g., glycer aldehydes and dihydr oxyacet one.
• Tet r oses (e.g. er yt hr ose) ar e quit e r ar e. Er yt hr ose t akes par t in t he synt hesis of
lignin and ant hocyanin pigment s.
• Ribose, r ibulose, xylulose and ar abinoses ar e pent oses. Xyluloses and ar abinoses
polymer ise t o for m xylans and ar abans which ar e cell wall mat er ials.
• Glucose, fr uct ose, mannose, galact ose ar e h exoses . These ar e whit e, sweet -t ast ing,
cr yst alline and ext r emely, soluble in wat er .
• Gl u cos e is t he univer sal sugar . It is also known as d e xt r os e or gr a p e s u ga r or
cor n s u ga r .
• F r u c t os e is t he most common for m of sugar in fr uit . It is also known as l e vu l os e. I t
i s t h e s we e t e s t a mon g n a t u r a l l y occu r r i n g s u ga r s .
• Monosacchar ides have ‘fr ee’ aldehyde or ket one gr oup which can r educe Cu
++
t o Cu
+
.
Hence, t hese ar e also called r educing sugar s.
2. Derived monosaccharides
• De oxy s u ga r – Loss of oxygen at om fr om r ibose yields deoxyr ibose, a const it uent of
DNA.
80 CSIR-NET Life Sciences
• Am i n o s u g a r – Mon os a cch a r ides h a vin g a n a min o gr ou p e.g., glu cos a min e,
galact osamine.
• Su ga r a ci d – e.g., Ascor bic acid, glucur onic acid, galact ur onic acid.
• Su ga r a l coh ol – e.g. glycer ol and mannit ol (pr esent in br own algae).
3. Oligosaccharides
The sugar s wit h limit ed number s (2–10) of monosacchar ides ar e called oli gos a cch a r i d e s .
These include t r isacchar ides, t et r a sacchar ides, hexasacchar ides, hept asacchar ides, et c.
• Di s a cch a r i d e s : These a r e for med by condensa t ion r ea ct ions bet ween t wo
mon os a cch a r i d e s (u s u a l l y h e xos e s ). Th e bon d for me d be t we e n t wo
monosacchar ides is called a gl ycos i d i c b on d . It nor mally for ms bet ween C-
at oms 1 and 4 of neighbour ing unit s (1,4 bond). Once linked, t he monosacchar ide
unit s ar e called r esidues.
(a) Ma l t os e (glucose + glucose), l a ct os e (glucose + galact ose), sucr ose (glucose
+ fr uct ose) ar e most common disacchar ides. Sucr ose is most abundant in
plant s and is known as ca n e s u ga r or t a b le s u ga r . It is t he sugar we buy
fr om ma r ket .
(b) On h y d r o l y s i s , di s a cch a r i des r el ea s e t h ei r r es pect i ve con s t i t u en t
monosacchar ides (e.g., hydr olysis of sucr ose yields one molecule each of
glucose and fr uct ose).
• Tr i s a c c h a r i d e s : Su ga r s composed of 3 mon osa cch a r ide u n it s a r e ca lled
t r i s a cch a r i d e s (e .g., r a ffi n os e ). Raffinose is a common sacchar ide found in
plant s. Upon hydr olysis, it yields one molecule each of glucose, fr uct ose and
galact ose.
• La r ge r ol i gos a cch a r i d e s a r e a t t a ch e d t o t h e ce l l me mb r a n e a n d ce l l
r e cogn i t i on i s d u e t o t h e i r p r e s e n ce . Th e y a l s o t a k e p a r t i n ant igen
specificit y.
Polymer isat ion of a lar ge number of small molecules r esult s in t he for mat ion of lar ge
molecules of high molecular weight s, which may be br anched or unbr anched. These ar e called
ma cr omole cu le s . These include some polysacchar ides, pr ot eins and nucleic acids.
Polysaccharides
These ar e polymer s of monosacchar ides and ar e br anced or unbr anched linear molecular chains.
These ar e insoluble car bohydr at es and ar e consider ed t o be non-sugar s. St ar ch, glycogen,
cellulose, pect in, hemicellulose, inulin and polysacchar ides.
P olys a cch a r i d e s ar e of t wo t ypes –
• Homop olys a cch a r i d e s – consist s of only one t ype of monosacchar ide monomer e.g.
st ar ch, glycogen and celluse, fr uct an, xylan, ar aban, galact an.
• He t e r op olysa cch a r i d e – consist s of mor e t han one t ype of monosacchar ide monomer
e.g. chit in, agar , ar abanogalact ans, ar abanoxylan et c.
Polysacchar ides ar e of t hr ee main t ypes – st or age (e.g. st ar ch and glycogen), s t r u c t u r a l
(e.g. chit in, cellulose) and mu cop olysa cch a r i d e s (e.g. ker at in sulphat e, chondr oit in sulphat e,
hyalur onic acid, agar , aliginic acid, car r agenin & hepar in).
Biochemistry 81
1. Storage polysaccharide
(i ) Starch, glycogen and i n u l i n ar e r eser ve food mat er ials. St ar ch t ur ns blue wit h
I od i n e.
• St ar ch is a polymer of glucose. It is t he major r eser ve food in plant s. St ar ch has
t wo component s – amylase (an unbr anched polymer ) and amylopect in (a br anched
polymer ).
• Amyl op e ct i n – consist s of 2000 – 200,000 glucose molecules for ming st r aight
chain and br anches (aft er 25 glucose unit s). Br anching point has a, 1-6 glycosidic
linkage.
• Amyl os e – consist s of a, 1-4 glycosidic linkage bet ween a-D glucose molecules.
It is st r aight chain of 200-1000 glucose unit s. It is helical; each t ur n consist s of 6
glucose unit s.
St ar ch molecules accumulat e in t he for m of layer s (st r at ificat ions) ar ound a shift ing or ganic
cent r e (hilum) t o for m st ar ch gr ains. In e cce n t r i c st ar ch gr ains, hilum lies on one side. These
ar e found in pot at oes. In con ce n t r i c st ar ch gr ains, hilum is pr esent in t he cent r e. These ar e
found in wheat , maize, and pea.
• Du mb -b e ll s h a p e d st ar ch gr ains ar e found in t he lat ex of Euphorbia.
• St ar ch gr ains wit h single hilum ar e called simple (e.g. maize); but t hose wit h
mor e t han one hilum ar e called compound (e.g. pot at o, r ice).
(ii) Glycogen : Glycogen is t he animal equivalent of st ar ch; many fungi also st or e it .
Glycogen t ur ns r ed-violet wit h iodine. It consist s of 30,000 glucose unit s joined by a,
1-4 bonds, much mor e banched t han st ar ch. Br anching point has a, 1-6 linkage,
br anching occur s aft er 10-14 glucose unit s.
(iii) I n u l i n : It is an unusual polysacchar ide and polymer of fr uct ose. It is used as a fr uct ose,
par t icular ly in r oot s and t uber s of t he family Composit ae (e.g. Dahlia t uber s).
II. Structure of polysaccharide
(a) Ce l l u l os e: Cellulose is t he main st r uct ur al polysacchar ide of plant s. It is t he most
abundant molecule on ear t h. It is almost confined t o plant s, t hough it is found in
pr imit ive fungi and lower inver t ebr at es also. It is st r uct ur al component of all plant
cell walls, const it ut ing, on an aver age, about 20-40% of t he wall. Cot t on fibr es cont ain
t he lar gest pr opor t ion (90%) of cellulose among nat ur al mat er ials.
Molecules ar e unbr anched, consist ing of 6000 – b – D glucose unit s joined by b, 1-4
linkages, and 2000 cellulose molecules fr om mi cr ofi b r i l . Rayon and cellophane ar e
similar t o ce l l u l os e xa n t h a t e. Cellulose acet at e is used t o pr epar e t r icot , double
knit , wr inkle pr oof and mout h pr oof clot hing, cigar et t e filt er s. Cellulose nit r at e is
used in pr opellent explosives. Car boxy met hyl cellulose is used as emulsifier and
smoot hing r eagent .
(b) Ch i t i n : Chit in is a polymer of acet yglucosamine (b, 1-4 glycosidic linkage) for ming
bundles of long par allel chains like cellulose. It occur s in t he cell walls of fungi (and
for ms exoskelet on in some animals especially ar t hr opods).
(c) P e ct i n a n d h e mi ce llu los e: Pect in and hemicelluloses ar e st r uct ur al polysacchar ides.
Pect ins ar e made up of a r a b i n os e , ga l a ct os e and ga l a ct u r on i c a ci d . Pect ic acid is
a homopolymer of t he met hyl est er of D-galact ur onic acid.
82 CSIR-NET Life Sciences
Mi d d l e l a me l l a which binds t he cells t oget her is composed of calcium pect at e. Due
t o t his subst ance, wat er absor pt ion capacit y of wall is incr eased.
Fr uit walls cont ain high per cent age of pect in. Dur ing r ipening, t he pect ins br eak
down int o sugar s r esult ing in t he sweet ning and loosening of fr uit s.
He mi ce llu los e s ar e homopolymer s of D-xylose linked by b 1-4, Xylans, ar abans,
galact ans ar e hemicelluloses. These ar e r ar ely used as food (e.g. dat es – Phoenix ).
III. Mucopolysaccharides
These ar e gelat inuous polysacchar ides for med fr om galact ose and mannose. Slimy subst ances
of bh i n d i , a ga r a ga r , a l gi n i c a ci d a n d ca r r a ge e n i n obt a i n e d fr om s e a we e d s a r e
mucopolysacchar ides. Mycopolysacchar ides ar e found in t he cell walls of bact er ia also.
(a ) Ke r a t i n s u l p h a t e – consist s of acet yl glucosamine, glaact ose and sulphur ic
acid, pr ovides st r engt h and flexibilit y t o skin and cor nea.
(b) Ch on d r oi t i n s u l p h a t e – consist s of flycur onic acid and acet yl glucosamine,
pr esent in t he vit r eous humor of eye, synovial fluid and cer ebr ospinal fluid et c.
(c) Hep a r in is a polymer of a, 1-4 glucosamine and glucur onic acid. It is ant icoagulant
pr esent in human blood. Husk of Plantago ovata and mucilage of Aloe barbedense
ar e medicinally used. Agar , alginic acid car r agenin ar e obt ained fr om mar ine
Br own algae (Phaeophyceae).
SUGARS
I. Sugar chemistry
A. Sugar s ar e car bohydr at es (Poly hydr oxy aldehydes or Ket ones)
1. empir ical chemical for mula (CH
2
O)
n
2. polyalcohols
3. hydr ophilic, polar
B. Car bonyl classificat ion of sugar s
1. Aldoses – aldehyde at t he C-1 car bon
2. Ket oses – car bonyl somewher e ot her t han C-1 (usually C-2)
C. St er eochemist r y of sugar s
1. all sugar s have at least one chir al car bon (except DHA)
2. nat ur ally occur ing sugar s ar e in t he D-for m
3. Fisher pr oject ion 2-D r epr esent at ion of st er ochem.
4. Epimer – sugar s differ ing in config. at one chir al car bon
D. Int r amolecular cyclizat ion of sugar s
1. Sugar s r eadily for m r ing st r uct ur es
(a) Condensat ion of a hydr oxyl wit h a car bonyl
(i) hemiacet al – cyclizat ion bet ween aldehyde and hydr oxyl
(ii) hemiket al – cyclizat ion bet ween ket one and hydr oxyl
(b) Differ ent size cyclic st r uct ur es may be for med
(i) pyr anose – six member r ing
(ii) fur anose – five member r ing
Biochemistry 83
2. Anomer ic car bon – new chir al cent er for med in t he cyclizat ion or for m many
esenzymes have specificit y for t he anomer ic car bon
3. Hawor t h pr oject ion – 3-D r epr esent at ion of cyclic st r uct ur e
E. Sugar chemist r y
1. Reducing sugar s
(a) car bonyl gr oup has r educt ive pot ent ial
(b) anomer ic car bon must be fr ee (not polymer ized)
(c) can r educe Cu
2+
(cupr ic) t o Cu
+
(cupr ous)
GLYCOLYSIS
1. All life pr ocesses r equir e ener gy. Much of t he food obt ained by aut ot r ophic and
het er ot r ophic nut r it ion is used as a sour ce of t his ener gy. The ener gy r ich compounds
ar e br oken down t o r elease ener gy by t he pr ocess of r espir at ion. Re s p i r a t i on i s a
p r oce s s b y wh i ch t h e e n e r gy of t h e food i s ma d e a va i l a b l e t o t h e ce l l s .
2. Glucose is t he most common r espir at or y subst r at e.
3. Phot osynt hesis is 10 t imes fast er t han r espir at ion.
4. About 50% of sugar for med in phot osynt hesis is used in r espir at ion.
5. Efficiency of r espir at ion is 40-45%.
6. Cellular r espir at ion is essent ially (a) a cat abolic pr ocess, (b) involves biological oxidat ion
of or ganic molecules and (c) r esult s in t he r elease of t he ener gy in t he for m of ATP.
ATP is a der ivat ive of AMP (Adenosine monophosphat e) t o which t wo addit ional
phosphat e gr oups ar e at t ached t hr ough an anhydr ide linkage. The t wo bonds ar e
indicat ed by symbol (~). When last high ener gy phosphat e bond is hydr olysed 8.9
k ca l ener gy is r eleased. When second phosphat e bond is br oken, 6.5 k ca lor ies of
ener gy is r eleased. However , if t he t hir d phosphat e gr oup (i.e., of AMP) is hydr olysed;
only 3.4 k ca l of ener gy is liber at ed. The ener gy r eleased by t he br eaking down of
phosphat e gr oups in ATP and ADP (Adenosine diphosphat e) is much mor e t han t he
ener gy r eleased on t he hydr olysis of most of t he ot her covalent bonds.
7. ATP is an inst ant sour ce of ener gy wit hin t he cell. It is mobile and t r anspor t s chemical
ener gy t o ener gy r equir ing pr ocesses wit hin t he cell. The hydr olysis of ATP r eleases
ener gy. Since ATP is found in all living cells, it is called u n i ve r sa l e n e r gy ca r r i e r
or “e n e r gy cu r r e n cy” of ce lls .
8. A r espir at or y subst r at e is a “subst ance which is oxidized t o yield t he ener gy necessar y
for cell maint enance and gr owt h”. These, include (i) car bohydr at es, (ii) fat s, (iii) pr ot eins.
9. Typ e s of r e s p i r a t i on :
(a) Ae r ob i c r e s p i r a t i on . It t a kes pla ce in t he pr esence of oxygen. Complet e
oxidat ion of t he subst r at e r esult s in t he for mat ion of CO
2
and wat er accompanied
by t he r elease of ener gy e.g.
C
6
H
12
O
6
+ 6O
2
→ 6CO
2
+ 6H
2
O + ener gy
(Glucose) (686 kcal)
In most for ms of life, r espir at ion occur s aer obically.
(b) An a e r ob i c r e s p i r a t i on . It t akes place in t he absence of oxygen and r esult s in
incomplet e degr adat ion of t he subst r at e. CO
2
and or ganic compounds like et hyl
84 CSIR-NET Life Sciences
alcohol, lact ic acid et c., ar e pr oduced accompanied by t he r elease of some ener gy,
wat er is not a pr oduct of t his r eact ion.
C
6
H
12
O
6
® 2C
2
H
5
OH + 2CO
2
+ ener gy
Usually, anaer obic r espir at ion occur s in t he deep seat ed t issues of plant s and
animals, in ger minat ing seeds, in fr uit s and among many micr oor ganisms e.g.
yeast s and bact er ia.
10. A lar ge var iet y of or ganisms (anaer obic) employ anaer obic r espir at ion as t heir major
ener gy yielding pr ocess. In fact , some bact er ia ar e even killed if exposed t o subst ant ial
amount of oxygen. These ar e t er med as obligat e anaer obes. The or ganisms which
can r espir e bot h in t he pr esence and t he absence of t he oxygen ar e called facult at ive
anaer obes.
11. Me ch a n i s m of r e s p i r a t i on : In aer obic as well as anaer obic r espir at ion, init ial
sequence of event s is t he same; collect ively t er med as glycolysis. Glycolysis (split t ing
of sugar ) r esult s in t he br eakdown of one molecule of glucose int o t wo molecules of
p y r u v i c a c i d. I t is complet ed in t he cyt opla sm of t he living cell not in t he
mit ochondr ia and does not r equir e t he pr esence of oxygen. It is also known as Embden -
Meyer h of-P a r n a s pat hway aft er t he names of t hr ee Ger man scient ist s who discover ed
it .
12. T h e p r o c e s s o f g l y c o l y s i s m a y b e s u b -d i v i d e d i n t o t h r e e s t e p s :
(a) Phosphor ylat ion, (b) Split t ing of fr uct ose-1, 6-diphosphat e, (c) For mat ion of pyr uvic
acid.
13. F a t e of p yr u vi c a ci d i n a e r ob i c r e s p i r a t i on : In aer obic r espir at ion, each pyr uvic
acid molecules ent er s a mit ochondr ion wher e it s oxidat ion is complet ed. It is car r ied
out in t he following phases: (a) For mat ion of acet yl coenzyme A and (b) Kr ebs cycle.
14. Acet yl CoA is pr oduced not only fr om pyr uvic acid, but also pr oduced as a r esult of â-
oxidat ion of fat t y acids under goes TCA cycle. Enzymes r equir ed for Kr eb’s cycle ar e
found in t he mat r ix of mit ochondr ia except succinat e dehydr ogenase which is bound
t o inner membr ane.
15. TCA: An a mp h i b oli c p a t h wa y: The TCA cycle is a cent r al met abolic pat hway playing
an impor t ant r ole in bot h: cat abolism and anabolism. It ser ves as a pat hway for
oxidat ion of not only car bohydr at es, but fat and pr ot eins also, ser ving as a common
oxidat ive pat hway (cat abolism). On t he ot her hand, t he int er mediat es of t he TCA
cycle ar e used in t he synt hesis of macr omolecules (anabolism).
• The t er m amphibolic (dual pur pose) is pr efer r ed for such pat hways.
16. One molecule of ATP (via GTP), four NADH, 1 FADH
2
and t wo molecules of CO
2
ar e
r eleased per molecule of pyr uvic acid oxidized. However , as t wo molecules of pyr uvic
acid ar e for med fr om one glucose molecule, TCA cycle must occur t wice for each
molecule of glucose r espir ed. Ther efor e, 2 ATP, 8 NADH and 2 FADH
2
a r e for med
fr om 2 molecules of pyr uvic acid (coming fr om one molecule of glucose).
17. Th e h ydr ogen ion s a n d elect r on s r emoved fr om r espir a t or y in t er media t es by
dehydr ogenat ion dur ing glycolysis and TCA cycle ar e ult imat ely oxidized t o wat er by
molecular oxygen. They r educe accept or molecules such as NAD
+
and FAD t o NADH
and FADH
2
r espect ively. Fr om NADH or FADH
2
,

t hese ar e passed along a chain of
int er mediat e subst ances (a chain of elect r on accept or s and t r ansfer molecules). The
enzymes necessar y for t he t r ansfer of elect r ons ar e pr esent in t he inner mit ochondr ial
Biochemistry 85
membr ane in an or der ed manner and also funct ion in a specific sequence. This
assembly of elect r on t r anspor t ing enzymes is called mi t och on d r i a l r e s p i r a t or y
ch a i n or t he e l e ct r on t r a n s p or t ch a i n .
Such mit ochondr ial membr ane is imper meable t o pr ot ons; t hese can not diffuse back
int o t he mat r ix acr oss t he membr ane. However , t hese can ent er t he membr ane via a
pr ot on channel est ablished by t he membr ane bound adenosine t r iphosphat ase (ATPase).
ATPase is a mult ienzyme complex cont aining t wo par t s F
0
and F
1
. F
1
component sit s
on t he sur face of t he membr ane while F
0
component is embedded in it for ming a
channel acr oss t he membr ane. Thr ough t his channel, t he pr ot ons flow t o F
1
. The
back of pr ot ons down t he pr ot on gr adient pr ovides ener gy for t he synt hesis of ATP
molecules. F
1
(act ing as ATP synt hesis) cat alyses t he synt hesis of ATP fr om ADP.
F or e a ch p a i r of p r ot on s fl owi n g b a ck i n t o t h e ma t r i x, on e mol e cu l e of ATP
i s syn t h esi zed . Thus, t he t hr ee pair s of pr ot ons, t hr e molecules of ATP ar e gener at ed.
Since FADH
2
denot es it s elect r ons t o CoQ and not t o FMN, only 4 pr ot ons ar e
t r anspor t ed out side t he membr ane. Ther efor e, t he backflow of t hese 4 pr ot ons t hr ough
F
0
-F
1
complex r esult s in t he for mat ion of only 2 ATP molecules.
18. Not yield of ATP in aer obic r espir at ion of glucose:
(a) In glycolysis, 4 molecules of ATP ar e pr oduced while t wo ar e consumed. Thus,
t her e is a dir ect gain of 2 ATP molecules in glycolysis.
(b) In TCA cycle, one molecule of ATP is pr oduced (via GTP) per molecule of pyr uvic
acid. Since 2 molecules of pyr uvic acid ar e pr oduced fr om one molecule of glucose,
2 molecules of ATP per glucose molecules ar e obt ained in TCA cycle.
• Thus, by dir ect phosphor ylat ion t her e is a gain of 4 ATP molecules in
r espir at ion.
(c) 10 NADH molecules ar e also pr oduced (2 in glycolysis and 8 fr om pyr uvic acid.
Each NADH molecule, t hr ough elect r on t r ansfer r eleases ener gy for t he synt hesis
of 3 ATP molecules. Thus, 30 mor e molecules of ATP ar e obt ained.
(d) Also, 2 molecules of FADH
2
ar e pr oduced in TCA cycle which yield 2 x 2 i.e. 4
ATP molecules.
(e) Thus, 4 + 30 + 4 = 38, i.e. a t ot al of 38 ATP molecules ar e yielded dur ing aer obic
r espir at ion of glucose.
(f) In most eukar yot ic cells, 2 ATP molecules ar e used up in t r anspor t ing 2 NADH
molecules pr oduced in glycolysis t o t he mit ochondr ion, in such cells, t her e in a
net gain of 36 ATP molecules inst ead of 38.
19. Though glycolysis and Kr ebs cycles const it ut e a major r espir at or y pat hway, ot her
r espir at or y pat hways also exist .
(a) P e n t os e p h os p h a t e p a t h wa y (Hexose mono-phosphat e shunt ) occur r ing in
cyt opla sm, r e q u i r e s oxyge n a n d i s a ma j or s ou r c e of 5-C s u ga r s . I t
p r od u ce d 36 ATP mole cu le s a n d ma y op e r a t e s i mu lt a n e ou s ly a lon g wi t h
n or ma l gl ycol yt i c p a t h wa y. Ne t yi e l d i s 35 ATP mol e cu l e s .
(b) Anot her cycle, Glyoxylat e cycle occur s in seeds t hat possess t issues r ich in fat
and enable st or ed fat t o be conver t ed int o car bohydr at es. Enzymes for t he cycle
ar e pr esent in glyoxysomes.
86 CSIR-NET Life Sciences
20. The accumulat ion of lact ic acid in t he muscle causes fat igue. However , dur ng r est ,
t he lact ic acid is again conver t ed int o pyr uvic acid and under goes aer obic r espir at ion.
21. Remember
(a) 264 Gms of CO
2
is liber at ed dur ing complet e oxidat ion of 180 gms of glucose.
(b) a-ket oglut ar ic acid is fi r s t d i ca r b oxyl i c a ci d for med dur ing Kr eb’s cycle.
(c) Number of ATP molecules for med by complet e oxidat ion of pyr uvic acid is 15.
(d) Me r cu r y i s u s e d i n a n a e r ob i c r e s p i r a t i on e xp e r i me n t s b e ca u s e i t d oe s
n ot r e a ct wi t h CO
2
.
(e) Rat e of r espir at ion is measur ed by r e s p i r ome t e r .
(f) F e r me n t a t i on was discover ed by Ga y Lu s s a c.
(g) Enzyme ext r act ed fr om yeast s which br ings about t he fer ment at ion is zymase.
(h) Fr uit s and seeds ar e st or ed at low t emper at ur e t o r educe t he r at e of r espir at ion.
(i) If a leaf is kept in sugar solut ion, t he r at e of r espir at ion incr eases.
(j) Di n i t r op h e n ol inhibit s ATP synt hesis (uncoupler of oxidat ive phosphor ylat ion).
(k) Ol i gomyci n inhibit s oxidat ive phosphor ylat ion (ener gy t r ansfer inhibit or ).
(l) Abnor mal r ise in r espir at or y r at e of r ipening fr uit s is called cl i ma ct r i c e .g.
b a n a n a .
22. F e r me n t a t i on d i ffe r s fr om a n a e r ob i c r e s p i r a t i on b y followi n g fe a t u r e s
(a) Fer ment at ion is an ext r acellular pr ocess.
(b) It usually occur s in t he pr esence of micr obes.
(c) Enzymes ext r act ed fr om cell can per for m fer ment at ion.
(d) A small quant it y of oxygen r at her st imulat es fer ment at ion, while anaer obic
r espir at ion occur s in t he absence of oxygen.
23. Higher concent r at ion of CO
2
and absence of O
2
adver sely affect t he r at e of r espir at ion
(except in anaer obic r espir at ion).
24. 1 molecule of glucose or fr uct ose pr oduces a t ot al of 38 ATP molecules out of which
t wo ar e used and t her efor e, net gain of ATP is 36. Out of t his 38 ATP, 4 ATP ar e
for med by dir ect (subst r at e) phosphor ylat ion and 32 ATP by oxidat ive phosphor ylat ion
t hr ough ETS and 2 by t hr ough GTP.
25. 1 molecule of NADH
2
for ms 3 ATP and 1 FADH2 for ms 2 ATP t hr ough ETS locat ed
on inner membr ane of mit ochondr ion.
26. One t ur n of Kr ebs cycle pr oduces 12 ATP, 11 ATP t hr ough ETS and 1 ATP by subst r at e
phosphor ylat ion.
27. ATP for mat ion is an ender gonic pr ocess and occur s in chlor oplast and mit ochondr ia.
28. RBC and muscles get ener gy by glycolysis.
29. Rat io of CO
2
for mat ion in aer obic and anaer obic r espir at ion is 3:1.
30. Rat io of ATP in aer obic and anaer obic r espir at ion is 18:1.
31. Kr eb’s cycle is basically a cat abolic cycle but also funct ions as anabolic cycle and
h en ce ca lled a mp h i b o l i c c y c l e. I t s st a r t ing pr oduct is cit r ic a cid which is a
t r icar boxylic acid. I t u n d e r goe s 4 oxi d a t i on s a n d 2 d e ca r b oxyl a t i on s t o pr oduce
CO
2
and H
2
O. It occur s in mat r ix of mit ochondr ia.
32. ETS, is locat ed on inner membr ane of mit ochondr ia, is r esponsible for ATP synt hesis
(oxidat ive phosphor ylat ion).
Biochemistry 87
33. ETS involves hydrogen and electron carriers and oxysom es (F
0
– F
1
par t icles) locat ed
on inner membr ane of mit ochondr ion. The sequence of t hese car r ier s is:
NADH
2
→ FMN → CoQ → Cyt b → Cyt c
1
→ Cyt c → Cyt a → Cyt a
3
.
34. Flow of elect r on in ETC is Fe
+3


Fe
+2


Fe
+3
.
35. Cyt ochr omes ar e Fe
+2
r ich int r insic pr ot eins. Cyt a
3

has bot h Cu
+2
and Fe
+2
and act s
as cyt ochr ome oxidase.
36. Rat io of t he volume of CO
2
liber at ed t o t he volume of O
2
absor bed dur ing r espir at ion
is called Respir at or y Quot ient , t he RQ.
(a) RQ is a n index of t ype of subst r a t e being r espir ed. Differ ent subst r a t es
(ca r bohydr a t es/fa t s/pr ot eins/or ga nic a cids) yield differ ent va lues of RQ on
oxidat ion.
(b) When car bohydr at es ar e complet ely oxidized, RQ = 1.
(c) When fat s and pr ot eins ar e r espir ed, RQ is less t han one.
(d) For or ganic acids (malic/oxalic acids et c.), RQ is gr eat er t han one.
(e) In succulent s (e.g., Opuntia, Bryophyllum) RQ = zer o as no CO
2
is r eleased (is
fixed int o or ganic acids).
(f) Dur ing anaer obic r espir at ion, RQ is infinit y (O
2
is not absor bed)
C
6
H
12
O
6
→ 6C
2
H
5
OH + 2CO
2
(RQ = 2 / 0 = ¥ )
37. Over all r eact ion
1. 2 pyr uvat es, 2 NADH and 2 ATP pr oduced per glucose
2. 2 st ages (hexose and t r iose st ages)
(a) hexose st age
(i) glucose phosphor ylat ed and cleaved t o for m t wo glycer aldehyde-3-P
(ii) uses 2 ATPs per glucose
(b) t r iose st age
(i) conver t s t wo glycer aldehyde-3-P t o t wo pyr uvat es
(ii) gener at es 4 ATPs and 2 NADH
3. Oxygen supply det er mines end pr oduct of pat hway
(a) Aer obic condit ions
(i) NADH is r eoxidized by ET/Ox Phos
(ii) pyr uvat e is used for acet yl CoA synt hesis
(b) Anaer obic condit ions
(i) NADH is r eoxidized by r educing pyr uvat e t o lact at e
(ii) lact ic acid is pr oduced
38. Pyr uvat e has mult iple fat es
1. Anaer obic glycolysis (higher or ganisms)
(a) Allows glycolysis t o cont inue wit hout O
2
(i) NAD
+
needs t o be r egener at ed fr om NADH
(ii) elect r on t r anspor t is inoper able
(iii) LDH r eoxidizes NADH
88 CSIR-NET Life Sciences
(b) Lact at e dehydr ogenase (LDH)
(i) Rever sible & r equir es NAD
+
(NADH)
(ii) lact ic acid pr oduced
(c) Physiology:
(i) skelet al muscle becomes anaer obic dur ing exer cise
(ii) lact ic acid is pr oduced so glycolysis can cont inue
(iii) dur ing r ecover y lact at e t r anspor t s t o t he liver for r econver sion t o
pyr uvat e or glucose
2. Anaer obic glycolysis (yeast )
(a) yeast gener at es et hanol in absence of O
2
(i) NAD
+
needs t o be r egener at ed fr om NADH; since elect r on t r anspor t
is inoper able
(iii) alcohol dehydr ogenase r eoxidizes NADH
(b) pyr uvat e decar boxylat ed t o acet aldehyde
(i) r equir es t hiamine pyr ophosphat e (TPP)
(ii) CO
2
causes bubbles in champagne and dough t o r ise
(c) t hiamine pyr ophosphat e
(i) der ived fr om t he vit amin t hiamine (B1)
(ii) involved in decar boxylat ion r eact ions
(d) acet aldehyde conver t ed t o et hanol
(i) cat alyzed by alcohol dehydr ogenase
(ii) NADH r eoxidized t o NAD
+
3. Aer obic oxidat ion
(a) pyr uvat e is oxidat ively decar boxylat ed t o for m acet yl CoA
(b) Acet yl CoA has many fat es including t he TCA cycle
39. Ther modynamic of glycolysis
A. Over all r eact ion: Fr ee ener gy change ( G) is highly negat ive
B. Individual r eact ions
1. React ions wit h posit ive Go’
(a) Fr ee ener gy changes ( G) is 0 or negat ive
(b) incr ease subst r at es and/or decr ease pr oduct s
(c) examples (i) aldolase
2. Highly exer gonic r eact ions ar e ir r ever sible
(a) hexokinase (b) phosphofr uct okinase (c) pyr uvat e kinase
40. Regulat ion of glycolysis
A. Hexose t r anspor t
1. Glucose t r anspor t ed int o cell by GLUT
- passive t r anspor t ; insulin causes mor e GLUT t o move t o membr ane
- insulin incr eases glucose upt ake by cells
2. GLUT is a family of glucose t r anspor t er s
(a) GLUT1: most t issues (not in liver or muscle)
(b) GLUT2: pancr eat ic – cells; r egulat e insulin r elease (?)
(c) GLUT4: hear t , skelet al muscle, and adipose t issue
Biochemistry 89
3. No GLUT in liver : upt ake of glucose by liver is not r egulat ed by insulin
B. Hexokinase, Glucokinase
1. Isozymes wit h differ ent pr oper t ies
Tissue Km Inhibition by Glucose-6-P
Hexokinase all 0.1 mM yes
Glucokinase liver, pancreas 10 mM no
C. Pyr uvat e kinase
1. 4 pyr uvat e kinase isozymes
(a) liver , kidney, and RBC isozymes ar e allost er ic; act ivat ed by Fr uct ose–1,6–
BP & inhibit ed by ATP
(b) liver and int est inal isozymes can be phosphor ylat ed
- phosphor ylat ion by pr ot ein kinase A; glucagon st imulat es pr ot ein kinase A
- phosphor ylat ed pyr uvat e kinase is less act ive; glucagon decr ease pyr uvat e
kinase act ivit y
D Phosphofr uct okinase-1
1. PFK-1 is allost er ically r egulat ed by sever al effect or s
2. ATP allost er ically incr eases Km for fr uct ose-6–P
- decr eases PFK act ivit y; AMP r elieves ATP effect
- ATP var ies lit t le (<10%), ADP and AMP var y gr eat ly
3. ADP act ivat es PFK-1 in mammals
4. Cit r at e inhibit s PFK-1
5. Fr uct ose 2,6-bisphosphat e (F-2,6-BP) is a pot ent act ivat or of PFK-
(a) One enzyme cont r ols synt hesis and degr adat ion of F-2,6-BP
(i) t wo enzymat ic funct ions on one pr ot ein
(ii) act ivit ies on separ at e domains
(iii) phosphor ylat ion of enzyme cont r ols act ivit y
- dephosphoenzyme synt hesizes F-2,6-BP
- phosphoenzyme conver t s F-2,6-BP t o F-6-P
- phosphor ylat ion r egulat ed by a cAMP-dependant kinase
- dephosphor ylat ion r egulat ed by a phosphat ase
(b) Phosphofr uct okinase-2 (PFK-2)
(i) synt hesizes F-2,6-BP fr om fr uct ose-6-P and ATP
(ii) inhibit ed by cit r at e & act ivat ed by Pi and indir ect ly by glucagon
(cAMP)
(c) Fr uct ose-2,6-bisphosphat ase
- hydr olyzes F-2,6-BP t o pr oduce fr uct ose-6-P
41. P a st eu r effect – P a st eu r Effect is a sudden change fr om anaer obic t o aer obic
r espir at ion due t o availabilit y of oxygen. The r at e of glucose consumpt ion is gr eat est
anaer obically
1. a er obic met a bolism of glucose pr oduces mor e ATP/glucose t ha n a na er obic
met abolism
90 CSIR-NET Life Sciences
2. ATP/ADP r a t io cont r ols glycolysis; glycolysis met a bolizes mor e glucose t o
gener at e equivalent wor d only ATP levels
AMINO ACIDS
I. Function of amino acids
A. Building blocks of polypept ides
1. polymer ized t o for m polypept ides
(a) linked by a pept ide bond
(b) synt hesized dur ing t r anslat ion of messenger RNA
2. pr imar y st r uct ur e of a pr ot ein is t he sequence of amino acids
3. bot h pept ides and polypept ides can be funct ional
St r uct ur e

OOC–CH
2
–CH
2
–NH
3
+
GABA
St r uct ur e
+
H
3
N–CH
2
–CH
2
–SO
3

TAURINE
B. Amino acids may be funct ional
1. neur ot r ansmit t er s
- glut amat e and aspar t at e (excit or y)
- glycine, t aur ine, and aminobut yr ic acid (GABA) (inhibit or y)
C. Pr ecur sor s t o ot her molecules
1. met abolic int er mediat es
- cit r ulline and or nit hine in ur ea cycle
- can be met abolized t o for m glucose or acet yl CoA
2. neur ot r ansmit t er s – ser ot onin, dopamine, epinephr ine, et c.
3. t yr osine (t hyr oid hor mone)
4. por phyr ins
5. cr eat ine (ener gy st or age)
6. hist amine (mediat or of immune r esponse)
7. nucleot ide synt hesis – S-adenosylmet hionine
II. Structure of amino acids
A. 20 st andar d alpha-amino acids
1. St r uct ur e.
- alpha-car bon
- alpha-amino gr oup
- alpha-car boxyl gr oup
- side chain (R gr oup)
2. “St andar d” amino acids ar e encoded by messenger RNA
3. Amino acids ar e abbr eviat ed by a 3-let t er and 1-let t er code
B. Some amino acids ar e not incor por at ed int o pr ot eins dur ing t r anslat ion
1. Modified amino acids
Biochemistry 91
(a) Hydr oxypr oline and hydr oxylysine
(i) hydr oxylat ed enzymat ically aft er t r anslat ion
(ii) impor t ant in collagen st r uct ur e (St r uct ur es of phosphoser ine, 4-OH-
pr oline, 5-OH-lysine)
(b) phosphoamino acids
(i) Tyr , Ser , Thr hydr oxyl gr oups can be phosphor ylat ed
(ii) impor t ant in act ivat ion and inhibit ion of enzymat ic or signalling act ivit y
2. Ot her impor t ant amino acids
(a) Ur ea cycle int er mediat es – or nit hine, cit r ulline, ar ginosuccinat e
(b) Thyr oid hor mone (t yr osine ® t hyr oxine)
Abbreviations for the 20 “standard” amino acids
alanine Ala A leucine Leu L
ar ganine Ar g R lysine Lys K
aspar agines Asn N met hioneine Met M
aspar t ic acid Asp D phenylalanine Phe F
cyst eine Cys C pr oline Pr o P
glycine Gly G ser ine Ser S
glut amine Gln Q t hr eonine Thr T
glut amic acid Glu E t r ypt ophan Tr p W
hist idine His H t yr osine Tyr Y
isoleucine Ile I valine Val V
III. Chemical Properties of Amino Acids
A. Physical characteristics
1. Char ge
(a) Amino acids ar e dipolar ions (zwit t er ions) at neut r al pH
(i) zwit t er ion is a dipolar molecule wit h pos. and neg. char ges spat ially separ at ed
(ii) definit ion of zwit t er ion in book is incor r ect
(b) Ionic st at es of amino acids depend on pH
(i) amino acids have t wo or t hr ee dissociable pr ot ons
(ii) pKa of t he dissociable pr ot on and t he pH det er mine it s degr ee of dissociat ion
H–H equation: pH = pKa + log{[A

]/[HA]}
2. Tit r at ion cur ve of an amino acid
(a) calculat ed using t he H-H equat ion
(b) Isoelect r ic point (pI) – pH at which t he molecule has a net char ge = 0 (aver age of
t he t wo appr opr iat e pKa values)
92 CSIR-NET Life Sciences
3. Polar it y
(a) nine nonpolar amino acids
(i) t end t o or ient t o t he inside of pr ot eins
(ii) Gly, Ala, Val, Leu, Ile, Met , Pr o, Phe, Tr p
(b) eleven polar amino acids
(i) t end t o or ient t o t he out side of pr ot eins
(ii) Ser , Thr , Tyr , Asp, Glu, Asn, Gln, Cys, Ar g, Lys, His
4. Hydr opat hicit y – index of solubilit y char act er ist ics in H
2
O
(a) combines hydr ophobic and hydr ophilic t endencies
(b) can be used t o pr edict pr ot ein st r uct ur e
Hydr ophobic Ile>Val>Leu>Phe>Met (list ed fr om most hydr ophobic)
Less Hydr ophobic Ala>Gly>Cys>Tr p>Tyr >Pr o>Thr >Ser
Hydr ophilic His>Glu>Asn>Gln>Asp>Lys>Ar g (ar g is most hydr ophilic)
5. UV absor bance
(a) ar omat ic a.a.s (Tr p, Tyr , Phe) absor b UV light
(b) absor bs UV light bet ween 260-280 nm
B. Stereochemistry
1. Most amino acids have opt ical act ivit y
(a) chir al cent er s ar e asymmet r ic cent er s (usually car bons)
(b) a-car bon of amino acids is chir al
(c) chir alit y yields st er eospecificit y
(d) Gly is not chir al (has no a-car bon)
2. L and D enant iomer s (st er eoisomer s)
(a) enant iomer s ar e nonsuper imposable mir r or images of t he same molecule
(i) L is levor ot at or y, D is dext r ar ot at or y
(b) L and D nomenclat ur e fr om L and D-glycer aldehyde (Fisher convent ion)
(i) not equivalent t o R and S
(ii) most nat ur al a.a.s ar e L
(iii) some a .a .s a r e R, some a r e S
(c) L and D for ms ar e chemically and physically dist inguishable
(i) differ ent act ivit y, melt ing point s, and spect r a
C. Cysteine can form disulfide bonds
1. cyst eine is t he r educed for m (sulfhydr yl)
2. cyst ine is t he oxidized for m (disulfide)
3. disulfide br idges for med bet ween cyst eines ar e impor t ant in pr ot ein st r uct ur e
Biochemistry 93
PROTEIN STRUCTURE AND PROTEIN FOLDING
I. Peptide bond – amide bond between alpha-amino and alpha-carboxyl groups
of 2 amino acids
A. Chemical properties
1. Pept ide bond is polar and planar
(a) elect r on r esonance st r uct ur e
(b) has par t ial (40%) double bond char act er
(c) amide gr oup is planar , usually t r ans
2. Synt hesis
(a) condensat ion pr oduces wat er
(b) ener gy r equir ed (ATP hydr olysis)
3. Pept ide bond is hydr olyzable
(a) acid hydr olysis gener at es fr ee amino acids
(i) 6N Hydr ochlor ic acid heat ed at 110°C for 24 hr in a vacuum
(b) base hydr olysis gener at es fr ee amino acids
(i) 4N Sodium hydr oxide heat ed at 100°C for 4 hr
(c) cyanogen br omide cleaves at t he COOH-t er minal side of Met
(d) enzymat ic hydr olysis of pept ide bonds by pr ot eases
(i) pept idases ar e specific for cer t ain amino acids
4. Polypept ides ar e polyampholyt es
(a) ampholyt e has bot h acidic and basic pKa values
(b) i s oe l e ct r i c p oi n t – pH at which t he net char ge is zer o
For example: H
3
N
+
–Ala–Lys–Ala–Ala–COO–
pKa of t he Alpha-car boxyl gr oup = 3.6 pKa of t he Alpha-amino gr oup = 8.0
pKa of t he delt a-amino of t he Lysine = 10.6
at pH = 1 t he net char ge is +2 at pH = 6 t he net char ge is +1
at pH = 14 t he net char ge is –1
t he isoelect r ic point pI = (pKa2 + pKa3)/2 = (8 +10.6)/2 = 9.3
B. Nomenclature
1. Size
(a) dipept ide (2 aminoacids & 1 pept ide bond), t r ipept ide (3 aminoacids & 2 pept ide
bonds)
(b) oligopept ide – sever al amino acids (up t o 20)
(c) polypept ides (mor e t han 20 amino acids). All pr ot eins ar e polypept ides
94 CSIR-NET Life Sciences
II. Physical Forces Governing Protein Conformation
A. Physical forces govern 3-D structure of proteins (Pauling and Corey)
1. bond lengt hs and angles should be dist or t ed as lit t le as possible
2. st r uct ur es must follow Van der Waal’s r ules for at omic r adii
3. pept ide bond is planar and t r ans
4. noncovalent bonding st abilizes st r uct ur e
5. confor mat ion can change wit hout br eaking bonds (flexibilit y)
B. Types of non-covalent forces important to protein conformation
1. Hydr ophobic for ces
(a) hydr ophobic r esidues or ient t o inside
(b) hydr ophilic r esidues or ient out
2. Van der Waal’s pot ent ial:
includes elect r on shell r epulsion, disper sion for ces, and elect r ost at ic int er act ions
3. Salt br idges, elect r ost at ic for ces
4. Hydr ogen bonds
C. Angles of rotation of the polypeptide chain determine structure
1. angles of r ot at ion ar ound alpha-car bon ar e [y (psi) and j (phi)]
(a) y (psi) is t he angle of t he alpha-car bon bond t o t he car bonyl-car bon
(b) j (phi) is t he angle of t he alpha-car bon bond t o t he amide-nit r ogen
2. pr imar y sequence and angles of r ot at ion for each alpha-car bon complet ely define
pr ot ein confor mat ion
3. only a small number of psi and phi angles ar e allowed
4. st at ist ical analysis of all pr ot eins yields gr oups of pr efer ed angles
(a) ar eas of r epeat ing (psi) and (phi) angles ar e secondar y st r uct ur es
III. Levels of Protein Structure
A. Primary Structure – amino acid sequence of a polypeptide
1. pr imar y st r uct ur e det er mines 3-dimensional st r uct ur e (Anfinsen)
2. always represented N H
2
-t er minus t o COOH-t er minus
B. Secondary structure – regular local conformation of linear segments of the polypeptide
chain
1. Secondar y st r uct ur e ar e st abilized by hydr ogen bonds bet ween amide and car bonyl
gr oups
2. Sever al t ypes of secondar y st r uct ur e
(a ) a l p h a -h e l i x
1. r ight handed helix
Biochemistry 95
2. 3.6 amino acids per t ur n, r ise per helix 5.4 Å, r ise per aminoacid 1.5 Å
3. car bonyl oxygen hydr ogen bonded t o 4
t h
amide hydr ogen (n®n+4)
4. amino acid R-gr oups or ient out
5. pr oline br eaks t he helix
(b) bet a -p lea t ed sh eet
(i) polypept ide chains side by side
(ii) polypept ide chains can be par allel or ant ipar allel
(iii) car bonyl oxygen hydr ogen bonded t o amide hydr ogen
(iv) bet a-st r and is a single pass of t he polypept ide
(c) r ever se t u r n , b et a -b en d
(i) allows a shar p t ur n in polypept ide chain
(ii) car bonyl oxygen hydr ogen bonded t o 3
rd
amide hydr ogen (n®n+3)
(iii) Glycine is r equir ed
3. Fibr ous pr ot eins demonst r at e secondar y st r uct ur e
(a) Fibr oin
(i) silk is fibr oin
(ii) ant ipar allel-bet a-pleat ed sheet
(b) alpha-Ker at in and t r opomyosin
(i) alpha-ker at ins in wool and hair and epider mal layer
(ii) t r opomyosin is a t hin filament in muscle
(iii) alpha-helix allows elast icit y
(iv) alpha-ker at in conver t s t o bet a-pleat ed sheet wit h heat or st r et ching
- disulfides ar e impor t ant t o maint enance of ker at in secondar y st r uct ur e
- alpha-ker at ins ar e bet a-pleat ed sheet s; feat her s and claws
(c) Collagen
(i) st r uct ur al pr ot ein; skin, bones
(ii) t r iple helix (not alpha-helix)
(iii) sequence (Gly-Xaa-Pr o) ‘ x or (Gly-Xaa-HyPr o) x
(iv) glycine r equir ed for t r iple helix t o for m, cont ains many modified amino
acids
(v) hydr oxypr oline st abilizes t he st r uct ur e, vit amin C r equir ed for hydr oxylat ion;
scur vy
C. Tertiary structure – overall folded conformation of the polypeptide
1. Physical for ces affect t er t iar y st r uct ur e
(a) Hydr ophobic for ces - hydr ophobic r esidues or ient t o inside & hydr ophilic or ient
out
(b) salt br idges, elect r ost at ic for ces
(c) Van der Waals r adii
(d) Hydr ogen bonds
(e) Disulfide br idges
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D. Quaternary structure – subunit structure
1. aggr egat ion of 2 or mor e subunit s
(a) het er o- or homo- polymer s
2. same for ces dr ive t er t iar y and quat er nar y st r uct ur e
E. Structural elements
1. Sequence mot if – small funct ional linear polypept ide sequence (may not be 3-D)
(a) signal pept ide
(b) ER-r et ent ion signal
(c) mit ochondr ial and nuclear t ar get ing signals
(d) RGD cell adhesion mot if
2. Super secondar y st r uct ur e (st r uct ur al mot if)
(a) smallest confor mat ional unit (may be funct ional)
(b) examples àà (helix-loop-helix), hair pin, bet a-bar r els
- Rossman fold, a nucleot ide binding sit e
- leucine zipper mediat es t r anscr ipt ion fact or dimer izat ion
- zinc finger is a DNA binding mot if
3. Domain
(a) t he par t of a polypept ide chain t hat can independent ly fold int o a t er t iar y st r uct ur e
(b) oft en domains have unit s of funct ion
(c) pr ot eins may cont ain one or many domains
IV. Protein folding
A. Folding occurs step-wise with several intermediates unfolded/secondary structure/
domains/molten globule/native tertiary structure
1. a collapsed st r uct ur e (molt en globule) occur s ver y quickly
2. st eps bet ween molt en globule and nat ive t er t iar y st r uct ur e usually occur slowly
(a) int er mediat es ar e isolat able
(b) mult iple pat hways ar e possible
B. Folding is driven by hydrophobic forces
C. Proteins can self assemble but in vivo folding is facilitated by proteins
1. Ch a p er on es ar e binding pr ot eins which assist folding
(a) chaper ones cause misfolded pr ot ein t o unfold r at her t han aggr egat e
(b) many chaper ones r equir e ATP hydr olysis for act ivit y
(c) mor e t han one chaper one may act simult aneously and sequent ially in t he folding
of a single pr ot ein
(d) chaper ones ar e specific for specific pr ot ein synt hesis pat hways (cyt osolic vs. mit o.
vs. endoplasmic r et iculum)
Biochemistry 97
2. Enzymes cat alyze kinet ically slow st eps infolding
(a) cis-t r ans pr olyl isomer ase
(i) bot h cis and t r ans pept ide bonds t o pr oline nat ur ally occur
(ii) t he isomer izat ion of pept idyl-pr oline bonds may be slow
(b) pr ot ein disulfide isomer ase
(i) cat alyzes disulfide bond for mat ion and isomer izat ion
D. Denaturation is unfolding
1. Requir es some input t o over come hydr ophobic for ces
(a) heat
(b) denat ur ant (ur ea or guanidinium)
2. Requir es r educt ant t o r educe disulfide br idges t o sulfhydr yls
V. Analytical Techniques in Protein Biochemistry
A. Determination of Amino Acid Composition
1. Amino acid analysis pr ovides % of each amino acid in pr ot ein
(a) Hydr olysis of polypept ide wit h 6N HCl
(b) Der ivit izat ion of amino acids wit h dansyl chlor ide PITC, or O-pht halaldehyde
(OPA)
(c) Liquid chr omat ogr aphic separ at ion of t he t agged amino acids
(d) Quant it at ion
2. Composit ion of a pr ot ein can be used t o ident ify a pr ot ein
B. Determination of primary sequence of a polypeptide
1. Pr epar at ion of pept ides for sequencing
(a) Removal of disulfide br idges
(i) r educing agent ( -mer capt oet hanol or dit hiot hr eit ol)
(ii) der ivat ize sulfhydr yls t o block disulfides fr om r eoxidizing
(b) Digest ion wit h cyanogen br omide
(i) CNBr cleaves at t he car boxyl side of met hionine r esidues
(c) Digest ion wit h pr ot eolyt ic enzymes
(i) use at least t wo differ ent enzymes
(ii) over lapping enzymes allows det er minat ion of pept ide sequence
(d) Separ at ion of pept ides
(i) pept ides separ at ed by chr omat ogr aphy
(ii) based on differ ences in ionic, polar , and/or hydr ophobic char act er ist ics
2. Ed ma n d e gr a d a t i on is used t o sequent ially det er mine aminoacid sequence
(a) PITC r eact s wit h t he N-t er minal amino acid
(b) St r ong acid cleaves t he pept ide bond bet ween t he 1st and 2nd amino acids
(c) Pr oduct is a PTH der ivat ive of amino acid #1
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(d) Det er mine ident it y of amino acid-PTH using HPLC chr omat ogr aphy
(e). Repeat st eps a-d
Not e: Edman degr adat ion has limit ed success wit h ver y long polypept ides
C. Determination of Molecular Mass
1. Gel Filt r at ion (Molecular exclusion chr omat ogr aphy)
(a) pr ot ein is loaded on a column of por ous beads
(b) small molecules can ent er t he beads, lar ge ones cannot
(c) an aqueous buffer moves t he pr ot ein t hr ough t he beads
2. SDS-polyacr ylamide gel elect r ophor esis (SDS-PAGE)
(a) pr ot ein is unfolded and coat ed wit h sodium dodecyl sulfat e (SDS) det er gent
(b) pr ot eins ar e loaded on an acr ylamide gel mat r ix
(c) elect r icit y moves t he pr ot eins t hr ough t he mat r ix
(d) low molecular weight pr ot eins move fast er (far t her )
(e) lar ge molecules migr at e fast er because t hey bypass beads
D. X-ray Crystallography and NMR
1. Physical t echniques t o ident ify 3-D st r uct ur e of a pur e pr ot ein
2. Requir es t r emendous t ime, effor t , and analyt ical r esour ces
1. To under st and t he major st r uct ur al feat ur es of immunoglobulins
2. To under st and t he int er act ion of immunoglobulins wit h ant igens
3. To lear n t he major classes of immunoglobulins
4. To under st and t he analyt ical uses of immunoglobulins
DNA STRUCTURE
A. DNA is a polymer of deoxyribonucleotide monophosphates
1. bases ar e adenine, guanine, cyt osine, and t hymine
2. linkage is t hr ough phosphodiest er bonds; 5' and 3' ends
3. ever y DNA has a specific sequence of nucleot ides—it s pr imar y st r uct ur e—genet ic
infor mat ion is st or ed in t he pr imar y st r uct ur e of DNA
B. Most DNA in a cell exists as the Watson-Crick double helix which is known as
B form DNA
Ma j or fe a t u r e s : r ight handed helix; bases on inside and sugar phosphat e backbone on out side;
base pair s ar e for med t hr ough hydr ogen bonding; A pair s wit h T; G pair s wit h C; bases ar e
per pendicular t o t he helical axis st acked on t op of each ot her and int er act ing t hr ough hydr ophobic
int er act ions and van der Waals int er act ions; 3.4 A per base pair ; ~10 base pair s per helical
t ur n; st r ands ar e ant ipar allel; st r ands ar e complement ar y
• A for m – having 11 base pair s (inst ead of 10 base pair s per t ur n), t he base pair s ar e
not per pendicular t o t he axis, but ar e t ilt ed.
Biochemistry 99
• C for m – like B for m, but having 9 base pair s per t ur n.
• D for m – like B for m, but have 8 base pair s per t ur n.
• Z-DNA–DNA wit h left handed coiling is called Z-DNA.
C. DNA supercoiling
1. a super coil is when t he double-helix t wist s ar ound it self
2. super coils ca n be posit ive or nega t ive but na t ur a l DNAs exist in t he nega t ive
super coiled for m
3. DNA can be super coiled if it is cir cular or if is linear and has fixed ends
4. super coiled DNA is mor e compact t han r elaxed DNA
5. negat ively super coiled DNA molecules ar e easier t o unwind t han r elaxed molecules
- DNA unwinding is r equir ed for r eplicat ion and t r anscr ipt ion
Top oi s ome r a s e s ar e enzymes t hat cat alyze changes in DNA super coiling
1. Type I t opoi som er a ses funct ion by br eaking a phosphodiest er bond of one st r and,
passing t he ot her st r and t hr ough t he br eak and r esealing t he br eak—t hey can only
r emove super coils
2. Type I I t opoi som er a ses funct ion by br eaking bot h st r ands and passing a double
st r and r egion t hr ough t he br eak befor e r esealing t he br eak—r equir e ATP
3. Topoisomer ases ar e t ar get s of numer ous chemot her apeut ic dr ugs: adr iamycin, VP16
(t enoposide), VM26 (et oposide), campt ot hecin
D. Nucleases
Enzymes t hat cat alyze hydr olysis of phosphodiest er bonds in nucleic acids
1. exonucleases cleave t er minal nucleot ides fr om eit her t he 5' or 3' end of a polynucleot ide
2. endonucleases cleave in t he int er ior of nucleic acid molecule—r est r ict ion enzymes
ar e endonucleases t hat cleave at specific sequences of DNA
E. Denaturation and renaturation of DNA
1. denat ur at ion is t he conver sion of t he double st r anded for m of DNA int o single st r anded
for m
(a) DNA can be denat ur ed by heat or alkaline t r eat ment
(b) The t emper at ur e at which half t he DNA is unwound is defined as t he melt ing
t emper at ur e (Tm)
–Tm is dependent on t he GC cont ent of t he DNA, on t he solvent , and on t he
ionic st r engt h
(c) Hyp e r ch r omi c e ffe ct : On denat ur at ion of dsDNA int o SS DNA UV absor pt ion
at 260 nm incr eases.
2. r enat ur at ion—under pr oper condit ions, complement ar y single-st r anded nucleic acids
can r enat ur e int o a double-st r anded for m
3. denat ur at ion/r enat ur at ion is t he basis of hybr idizat ion exper iment s—t his t ype of
analysis is cent r al t o r ecombinant DNA t echnology and gene manipulat ion
100 CSIR-NET Life Sciences
F. Packaging of DNA in eukaryotes
1. DNA is packaged in t he nucleus as a nucleopr ot ein complex called chr omat in
2. Levels of chr omat in packaging:
(a) Nu cl e os ome s : ~200 bp DNA and hist ones
(i) nucleosome cor e par t icles consist of ~140 bp of DNA wr apped ar ound a
pr ot ein oct amer consist ing of 2 subunit s each of hist ones H2A, H2B, H3,
and H4
(ii) linker DNA is t he DNA bet ween t wo cor e par t icles; hist one H1 binds t o t he
linker DNA and t he cor e par t icle
(b) 30 n m fi ber (Solen oi d ): nucleosomes ar e wound int o a solenoid-like st r uct ur e—
r equir es hist one H1 binding t o ever y linker DNA
3. chr omat in is t he t emplat e for r eplicat ion and t r anscr ipt ion and t he subst r at e for
DNA r epair and r ecombinat ion
DNA REPLICATION
A. DNA r eplicat ion is semiconser vat ive
B. DNA r eplicat ion is (usually) bidir ect ional
C. Chain gr owt h occur s by addit ion of deoxynucleot idyl monomer s t o t he end of a DNA
chain
1. DNA is always synt hesized in t he 5' t o 3' dir ect ion
2. This r eact ion is car r ied out by DNA polymer ases
(a) gener al r equir ement s of DNA polymer ases
(i) deoxynucleot ide t r iphosphat es (dATP, dGTP, dCTP, TTP)
(ii) DNA t emplat e
(iii) a pr imer chain wit h a fr ee 3'-OH—not e t hat DNA polymer ases cannot
st ar t wit h a single nucleot ide
D. At a r eplicat ion for k one of t he new st r ands is synt hesized in a cont inuous manner
(t he leading st r and) and t he ot her is synt hesized in a discont inuous manner (t he
lagging st r and)
E. Molecular event s of DNA r eplicat ion (E. coli chr omosomal r eplicat ion)
1. I n i t i a t i on
(a) an init iat or pr ot ein (dnaA pr ot ein in E. coli) binds t o an or igin of r eplicat ion
and “melt s” a shor t DNA sequence —an or igin is a unique sit e on t he
chr omosome wher e r eplicat ion begins, it consist s of binding sit es for t he
init iat or pr ot ein and a flanking AT-r ich sequence
(b) helicase (dnaB/dnaC complex) binds t o melt ed r egion and fur t her unwinds
par ent al st r ands
(c) SSB (single-st r and binding pr ot ein) binds t o unwound r egion t o pr event
r eannealing and st abilize t he single-st r anded for m at t he r eplicat ion for k
(d) an RNA pr imer is synt hesized by pr imase (pr imase is par t of a mult i-pr ot ein
complex called t he pr imosome)
(e) DNA polymer ase III (holoenzyme) is assembled
Biochemistry 101
2. El on ga t i on
(a) DNA polymer ase III ext ends RNA pr imer s
(b) helicase cont inues t o unwind par ent al DNA st r ands ahead of polymer ase
(i) t he leading st r and cont inues unint er r upt ed (DNA pol III has high
pr ocessivit y)
(ii) on t he lagging st r and, appr oximat ely 1000 bp ar e r eplicat ed befor e
pr imase must synt hesize a new RNA pr imer t o be elongat ed by DNA
pol III. —Okazaki fr agment s
3. DNA p ol yme r a s e I (i n E. col i )
Klenov fr a gment ha ve 5’®3’ polymer a se (fills in t h e ga p) a n d 3’®5’
Exonuclease act ivit y (Pr oofr eading act ivit y). Anot her unit r emoves RNA
pr imer (5'-3' exonuclease act ivit y) and
4. DNA l i ga s e
Seals t he nicks—E. coli DNA ligase r equir es NAD
+
and Bact er iophage l
r equir es ATP
5. Te r mi n a t i on of r eplicat ion t akes place wit hin a r egion of t he cir cular E. coli
chr omosome called t er
F. The r eplisome is a lar ge mult ipr ot ein “machine” t hat is t hought t o r eplicat e bot h t he
leading and lagging st r and simult aneously
G. Pr oofr eading: Pol III (and Pol I) has 3' —5' exonuclease act ivit y —if incor r ect base is
inser t ed it can back up 1 nucleot ide and t hen cont inue polymer izat ion
H. DNA r eplicat ion in eukar yot es
1. enzymat ically t he mechanism is basically t he same as in pr okar yot es
2. each chr omosome has mult iple or igins
3. DNA is r eplicat ed only dur ing S phase of t he cell cycle and only once dur ing each
cell cycle
4. t he subst r at e (t emplat e) for r eplicat ion is chr omat in
5. a special enzyme called t elomer ase is necessar y for r eplicat ing t he ends of linear
chr omosomes
(a) t he ends of eukar yot ic chr omosomes ar e called t elomer es and ar e made up
of shor t r epet it ive sequences
(b) t elomer ase is an enzyme t hat cont ains bot h pr ot ein and RNA component s
(c) t he RNA component is used as a t emplat e t o synt hesize new t elomer e r epeat s
(d) t elomer ase and cancer
DNA REPAIR
A. DNA damage caused by ultraviolet light
1. cyclobut ane-t ype pyr imidine dimer is t he major phot opr oduct for med
2. a second pr oduct , t he 6-4 phot opr oduct , is for med in about 10 % of UV induced
pyr imidine dimer s
3. t he cyclobut ane t ype dimer can be r ever sed by a pr ocess called phot or eact ivat ion
102 CSIR-NET Life Sciences
(a) t his is car r ied out by an enzyme called DNA phot olyase (phot or eact ivat ing
enzyme)
(b) impor t ance of t he phot olyase enzyme in humans is quest ionable
4. DNA phot opr oduct s can also be r epair ed by excision r epair
B. Spontaneous deamination of cytosine
1. deaminat ion of cyt osine is common and r esult s in t he conver sion of cyt osine t o ur acil
2. can be r epair ed by excision r epair pr ocess
(i) ur acil-DNA glycosylase hydr olyzes N-glycosidic bond t o r emove ur acil base
(ii) AP endonuclease r emoves deoxyr ibose-phosphat e
(iii) gap is ext ended by exonuclease
(iv) gap is filled by DNA polymer ase I and nick is sealed by DNA ligase
C. Depurination
Pat hway is similar t o t hat above except t hat excision r epair begins wit h AP endonuclease
D. DNA damaged by alkylating agents
1. Some simple alkylat ing agent s:
2. examples of pr oduct s of alkylat ing agent s:
3. many of t hese pr oduct s can be r epair ed by excision r epair t hat is init iat ed by specific
glycosylases
4. some damage r esult ing fr om met hylat ion can be r ever sed by met hylt r ansfer ases
(a) O6-met hylguanine-DNA met hylt r ansfer ase
E. Mismatch repair
1. mismat ches can occur when DNA polymer ase inser t s t he wr ong nucleot ide dur ing
r eplicat ion
2. mismat ch r epair is “coupled” t o r eplicat ion
3. How do t he mismat ch r epair enzymes dist inguish which base is incor r ect ?
(a) pa r ent a l DNA is met hyla t ed—in E. coli a n enzyme ca lled Da m met hyla se
met hylat es t he C in bot h st r ands at t he sequence GATC
(b) immediat ely aft er r eplicat ion only t he par ent al st r ands ar e met hylat ed (t he DNA
is hemimet hylat ed)
4. t he defect ive gene in one for m of her edit ar y colon cancer was r ecent ly found t o be
t he human homologue of mut S
F. Recombinational repair
1. occur s dur ing DNA r eplicat ion
2. major st eps:
(a) DNA polymer ase skips over damaged DNA leaving a gap opposit e t he lesion
(b) t he undamaged par ent al st r and r ecombines int o t he gap (t his is facilit at ed by
r ecA pr ot ein in E. coli)
(c) t he new gap in t he par ent al st r and is filled by DNA polymer ase and ligase
Biochemistry 103
G. Genetic Defects in DNA repair and human disease
1. Xer oder ma pigment osum is an inher it ed disease t hat is char act er ized by sever e
phot osensit ivit y and a ver y high incidence of skin cancer s. It is due t o defect ive
excision r epair .
2. Bloom’s syndr ome.
3. Cockayne’s syndr ome
4. Fanconi’s anemia
5. At axia t elangiect asia
RNA STRUCTURE, TRANSCRIPTION, RNA PROCESSING
I. Structure of RNA
A. RNA is a polymer of r ibonucleot ide monophosphat es
1. pur ine bases ar e adenine and guanine; pyr imidine bases ar e cyt osine and ur acil
2. RNA molecules can have ext ensive secondar y st r uct ur e
(a) int r amolecular base pair ing
(b) r egions of base pair ing in RNA for m an A-t ype double helix
(c) many secondar y st r uct ur es of RNA have defined funct ional r oles
II. Classes of RNA molecules
A. Messenger RNA (mRNA): t r anscr ibed by RNA polymer ase II in eukar yot es; encode
pr ot eins
B. Ribosomal RNA (r RNA)
1. 18 S, 28 S, and 5.8 S r RNAs ar e t r anscr ibed by RNA polymer ase I in eukar yot es
2. 5 S RNA is anot her t ype of RNA associat ed wit h r ibosomes but is t r anscr ibed by
RNA polymer ase III in eukar yot es
3. r RNA ser ve st r uct ur al and cat alyt ic r oles in r ibosomes
C. Tr ansfer RNA (t RNA): t r anscr ibed by RNA polymer ase III in eukar yot es.
Not e: all t hr ee t ypes of RNA’s above ar e t r anscr ibed by t he same RNA polymer ase in pr okar yot es
D. Numer ous ot her small RNA’s ar e also found in cells—in eukar yot ic cells t hese can be
put int o t wo gener al classes: snRNA = small nuclear RNA & scRNA = small cyt oplasmic
RNA
1. snRNA’s and scRNA’s ar e found complexed wit h pr ot eins and car r ied a var iet y of
cellular funct ions (snRNP and scRNP)
III. Transcription
A. RNA molecules ar e t r anscr ibed fr om a DNA t emplat e by RNA polymer ases
B. r equir ement s for RNA polymer ases: DNA t emplat e, ATP, GTP, CTP, UTP, Mg
++
(no
pr imer is necessar y for RNA polymer ase)
104 CSIR-NET Life Sciences
C. chain gr owt h is fr om 5' t o 3'
D. Gener al mechanism of t r anscr ipt ion
IV. RNA synthesis in bacteria
A. Steps of transcription
1. I n i t i a t i on
(a) RNA polymer ase binds t o DNA and migr at es t o t he pr omot er
(i) a pr omot er is a specific DNA sequence t hat cont ains a sit e for t r anscr ipt ional
init iat ion—E. coli pr omot er cont ain a -10 r egion and a -35 r egion t hat ar e
impor t ant in binding polymer ase
(ii) specific int er act ion bet ween t he pr omot er and RNA polymer ase r equir es
sigma fact or
(iii) init ia l complex bet ween polymer a se a nd pr omot er is ca lled t he closed
complex
(b) RNA polymer ase unwinds ~18 bp for ming an open complex
(c) fir st nucleot ide, which is almost always a pur ine, int er act s wit h t he open complex
by binding t o polymer ase and base pair ing wit h t he complement ar y nucleot ide
in t he t emplat e st r and
2. El on ga t i on
(a) t h e s econ d n u cl eot i de bi n ds t o t h e pol ymer a s e-t empl a t e compl ex a n d a
phosphodiest er bond is for med
(b) sigma fact or is r eleased and polymer ase moves down t he t emplat e, unwinding
t he t emplat e and cat alyzing t he addit ion of each successive nucleot ide
(c) appr oximat ely 12 nucleot ides of t he gr owing RNA chain ar e base pair ed wit h t he
DNA t emplat e dur ing t he elongat ion phase
3. Te r mi n a t i on
(a) Ter minat ion r equir es special t er minat ion signals
(i) r ho-independent t er minat ion
(ii) r ho-dependent t er minat ion
B. Inhibitors of bacterial RNA synthesis
1. act inomycin D—binds t o DNA by int er calat ing bet ween base pair s
2. r ifampicin (r ifamycin)—binds t o t he bet a subunit of bact er ial RNA polymer ase
- also inhibit s mit ochondr ial RNA polymer ase
- used t o t r eat t uber culosis which is r esist ant t o most ot her ant ibiot ics
V. DNA synthesis in eukaryotes
A. transcription and processing of mRNA
1. most t r anscr ipt ion unit s have exons and int r ons
2. Init iat ion of t r anscr ipt ion by RNA polymer ase II
(a) polymer ase II does not r ecognize a specific sequence in t he pr omot er ; it is
posit ioned at t he cor r ect sit e by int er act ion wit h t r anscr ipt ion fact or s
Biochemistry 105
(b) many genes cont ain a sequence called t he TATA box approximat ely 30 bp upst ream
fr om t he t r anscr ipt ional st ar t sit e; a t r anscr ipt ion fact or named TFIID specifically
binds t o t he TATA box t o help posit ion t he polymer ase at t he init iat ion sit e.
Some genes lack a TATA box but also ut ilize a sequence-specific t r anscr ipt ion
fact or t o t ar get polymer ase t o t he init iat ion sit e.
3. Capping of t he 5' end of t he t r anscr ipt
(a) ver y soon aft er t he t r anscr ipt is init iat ed a 7-met hylguanosine “cap” is added t o
t he 5' end of t he t r anscr ipt by guanosyl t r ansfer ase
(b) 7mG cap is an impor t ant signal for t he t r anslat ion pr ocess and may also help
pr ot ect t he message fr om degr adat ion
4. Polyadenylat ion by Poly (A) polymer ase
(a) t he 3' end of (most ) t r anscr ipt s is modified by polyadenylat ion
(b) t he poly A t ail is usually about 200-250 nucleot ides
(c) t he poly A t ail is not encoded by t he gene but is added post -t r anscr ipt ionally
(d) polyadenylat ion r equir es a specific sequence (AAUAAA) in t he mRNA
(e) no specific t er minat ion signal is known for eukar yot ic genes and t r anscr ipt ion
pr oceeds past t he polyadenylat ion signal
5. mRNA splicing
(a) int r on sequences ar e r emoved fr om pr imar y t r anscr ipt s by a pr ocess called splicing
(b) splicing r equir es consensus splicing signals at t he 5' and 3' ends of t he int r on, a
consensus br anch point 10-40 bases upst r eam of t he 3' end of t he int r on
(c) splicing is car r ied out by a lar ge complex called t he spliceosome—t he spliceosome
is assembled fr om small nuclear r ibonucleopr ot ein (snRNP’s)
B. Synthesis and processing of rRNA in eukaryotes
1. r RNA is synt hesized in t he nucleolus by RNA polymer ase I
2. genes for r RNA’s ar e found in mult iple copies in t andem ar r ays on t he chr omosome
3. all t hr ee major r ibosomal RNA’s (28S, 18S, 5.8S) ar e synt hesized as par t of t he same
pr ecur sor t r anscr ipt which is pr ocessed by a ser ies of cleavage st eps t o pr oduce t he
mat ur e r RNA’s
4. assembly of r RNA’s and r ibosomal pr ot eins int o lar ge and small r ibosomal subunit s
also t akes place in t he nucleolus
C. Synthesis and processing of tRNA
1. synt hesized by RNA polymer ase III in eukar yot es
2. pr imar y t r anscr ipt is a pr ecur sor t hat gener ally has ext r a nucleot ides on bot h t he 5'
and 3' ends; some t RNA genes also have int r ons but splicing is by a complet ely differ ent
mechanism t han wit h mRNA
3. t he ext r a nucleot ides ar e r emoved fr om t he ends and t hen 3 nucleot ides (CCA, t hese
ar e not encoded by t he gene) ar e added t o t he 3' end post -t r anscr ipt ionally
4. t he bases of t RNAs under go ext ensive post -t r anscr ipt ional modificat ion, up t o 10 % of
t he nucleot ides can be modified
5. mat ur e t RNAs have ext ensive secondar y and t er t iar y st r uct ur e t hat is impor t ant for
t heir funct ion
106 CSIR-NET Life Sciences
PROTEIN SYNTHESIS
I. Overview of Translation
A. Tr anslat ion is t he pr ocess by which a polypept ide chain is synt hesized by r ibosomes
using t he sequence of codons in an mRNA t o dir ect t he sequence of amino acids.
B. Tr anslat ion is t he ult imat e st ep in gene expr ession.
C. The ener gy cost for pr ot ein synt hesis is ver y high.
1. Only a small fr act ion of t he ener gy input of t r anslat ion is needed t o for m t he
pept ide bond.
2. The major it y of ener gy is invest ed t o assur e t hat t he sequence of t he polypept ide
is cor r ect .
3. If incor r ect polypept ides (e.g. enzymes) ar e made by t he cell, it could have
devast at ing effect s on cell funct ion.
D. The mRNA is always r ead fr om 5' t o 3'.
E. The polypept ide is always synt hesized in t he dir ect ion of amino-t er minus t o car boxyl-
t er minus.
II. Components of Translation
A. mRNA
The mRNA ser ves at t he t emplat e t hat will det er mine t he sequence of amino acids in t he new
polypept ide.
1. St r u ct u r e of mRNA
(a ) 5' u n t r a n s l a t e d r e gi on (5' UTR).
(i) This is t he sequence of t he mRNA ext ending fr om t he 5' end of t he mRNA
t o t he init iat ion codon.
(ii) It is not t r anslat ed int o polypept ide sequence.
(iii) It has a funct ion analogous t o t he funct ion of a pr omot er on a gene. It will
dir ect t he binding of t he r ibosome t o t he init iat ion codon.
(b) I n i t i a t i on cod on
(i) This is t he t r iplet codon at which polypept ide synt hesis begins.
(ii) It is always AUG and codes for a met hionine.
(iii) As a r esult , a ll polypept ides a r e synt hsized wit h a n a mino-t er mina l
met hionine.
(c) Cod i n g r e gi on
(i) This is t he sequence of mRNA t hat cont ains t he consecut ive t r iplet codons
t hat dir ect polypept ide synt hesis.
(ii) This r egion spans fr om t he st ar t codon t o t he st op codon.
(iii) The coding r egion is oft en r efer r ed t o as t he op e n r e a d i n g fr a me or ORF.
(d ) St op cod on
(i) This is t he t r iplet codon t hat signals t he t er minat ion of t r anslat ion.
(ii) Ther e ar e t hr ee possible st op codon sequences: UAA, UAG, UGA.
(iii) St op codons have no cor r esponding t RNA or amino acid.
Biochemistry 107
(e) 3' u n t r a n s la t e d r e gi on (3' UTR)
(i) This is t he nucleot ide sequence downst r eam fr om t he st op codon.
(ii) It ext ends fr om t he st op codon t o t he 3' end of t he mRNA
(iii) It does not code for amino acid sequence.
(iv) It may funct ion in st abilizing t he mRNA.
B. tRNA
1. The funct ion of t RNA is deliver t he cor r ect amino acid t o t he r ibosome as dir ect ed by
t he codon on t he mRNA for incor por at ion int o t he polypept ide.
2. The t RNA has t wo impor t ant funct ional feat ur es
(a) A t RNA is covalent ly bound t o an amino acid at it s 3' end.
(i) This for m of t RNA is called an a mi n o a cyl t RNA.
(ii) Each t RNA will be bound t o only one of t he t went y possible amino acids.
However , t her e can be mult iple t RNAs for each amino acid.
(b) The t RNA cont ains a t r iplet sequence of nucleot ides t hat ar e complement ar y t o
t he t r iplet codons of t he mRNA. This sequence is called t he a n t i -cod on .
(i) The covalent ly bound amino acid and t he ant i-codon st r ict ly cor r espond t o
one a not her .
(ii) As a r esult , t he pair ing of codon of t he mRNA and t he ant i-codon of t he
amino acyl-t RNA dur ing t r anslat ion will deliver t he cor r ect amino acid t o
t he gr owing polypept ide chain.
3. St r u ct u r e of t RNA
(a ) Cl ove r l e a f s t r u ct u r e
(i) 73-93 bases in lengt h
(ii) Many unusual bases (Psuedo Ur acil, Thymine, Dihdr oxy Ur acil et c.)
(iii) Th e n u cleot ides for m ma n y i n t r a -c h a i n b a s e p a i r s, r es u lt in g in a
s e con d a r y s t r u ct u r e t hat r esembles a clover leaf.
(b) Acce p t or St e m
(i) This is a r egion of t he t RNA t hat is t he sit e of at t achment for t he appr opr iat e
amino acid.
(ii) It is for med by seven r egular Wat son & Cr ick base pair s bet ween t he 5' and
3' ends of t he t RNA.
(iii) The 3' t er minal end of all t RNAs is always CCA-OH.
(a) It is not base pair ed and is t he sit e of at t achment of t he amino acid.
(b) The amino acid is covalent ly bound t hr ough an e s t e r l i n k a ge bet ween
t he car boxyl gr oup of t he amino acid and t he 3' hydr oxyl gr oup of t he
r ibose of t he t RNA.
(c) An t i cod on Loop
(i) The ant i-codon loop cont ains t he t hr ee nucleot ide sequence t hat is t he r ever se
complement of t he codon of mRNA t o which it cor r esponds.
(ii) It consist s of a t ot al of seven unpair ed bases; t hr ee of which ar e t he ant i-
codon.
4. Wob b le Hyp ot h e s i s
(a) Four common bases in mRNA can be ar r anged in 43 or 64 differ ent combinat ions
(b) How does t his r elat e t o t he 20 aa’s available for t r anslat ion?
108 CSIR-NET Life Sciences
(i) Genet ic code is r edundant
(a) 3 codons used as st ops UAA, UAG, UGA.
(b) 61 available as codons
(c) 1965, Fr a ncis Cr ick pr oposed t he Wobble Hypot hesis t o a ccount for t his
inconsist ency
(i) Codons in mRNA and ant i-codons int er act in an ant i-par allel manner at t he
r ibosome so t hat posit ions 1 and 2 of t he codon for m nor mal Wat son-Cr ick
base pair s.
(ii) The t hir d posit ion allows non-st andar d base pair ing or wob b le.
(d) Consequences: Differ ent ant i-codons can pair wit h one codon.
C. Ribosome
1. Per for ms 3 essent ial funct ions
(a) Select ion of mRNA init iat ion sit e.
(b) Ensur es cor r ect pair ing of codons and ant icodons.
(c) Cat alyzes for mat ion of pept ide bonds.
2. Ribosomes ar e lar ge pr ot ein-RNA complexes t hat ar e divided int o a lar ge and small
subunit .
(a) Lar ge subunit .
(i) In pr okar yot es t his subunit is called t he 50S subunit .
(ii) In eukar yot es t his subunit is called t he 60S subunit .
(iii) It cont ains t he act ive sit e for cat alyzing polypept ide bond for mat ion. This
enzyme wit hin t he r ibosome is called t he p e p t i d yl t r a n s fe r a s e.
(b) Small subunit
(i) In pr okar yot es t his subunit is called t he 30S subunit .
(ii) In eukar yot es t his subunit is called t he 40S subunit .
(iii) It is pr imar ily r esponsible for posit ioning t he r ibosome at t he int it iat ion
codon and pair ing codon and ant i-codon dur ing synt hesis.
III. Mechanism of Translation
A. Three steps in translation process
1. I n i t i a t i on – The small and lar ge subunit s of t he r ibosome bind at t he init iat ion codon
and t he met hionine-t RNA ant icodon pair s wit h t he st ar t codon.
2. El on ga t i on – The r ibosome pr oceeds down t he mRNA one t r iplet codon at a t ime,
posit ioning t he cor r ect amino acyl-t RNA wit h t he codon, and cat alyzing polypept ide
synt hesis.
3. Te r mi n a t i on – The r ibosome encount er s a st op codon. The polypept ide, t RNA, and
mRNA ar e r eleased. The small and lar ge subunit s dissociat e fr om one anot her .
B. Prokaryotic translation
1. I n i t i a t i on – t hr ee st eps t o init iat ion
(a) mRNA binds t o 30S.
(i) Ribosome aligned by base pair ing of a r egion of 16S r RNA of t he 30S
r ibosomal subunit t o a r egion on t he mRNA 6-10 bases upst r eam of t he
init iat ion codon.
Biochemistry 109
(ii) The region is called the Sh i n e /Da lga r n o se q u e n ce .
(b) Met hionyl-t RNA binds t o 30S-mRNA complex.
(i) Fir st aa-t RNA is posit ioned complement ar y t o t he fir st codon by t he 30S
r ibosomal subunit .
(ii) In pr okar yot es t he fir st amino acyl-t RNA is always for myl met hionyl t RNA
(fMet -t RNAfMet )
(iii) Th er efor e, a l l pr ot ei n s i n pr oka r yot es a r e s yn t h es i zed wi t h for myl
met hionine as t heir fir st amino acid.
(iv) This complex is called t he 30S pr e-init iat ion complex.
(c) 50S subunit binds t o 30S-t RNA-mRNA complex.
(i) This complex is called t he i n i t i a t i on comp l e x .
(ii) Two sit es for amino acyl-t RNA binding on 50S subunit .
(a) The p e p t i d yl s i t e or P -s i t e. This is t he sit e wher e t he gr owing
polypept ide will be anchor ed t o t he r ibosome. The polypept ide is held
in place by covalent at t achment of it s car boxyl-t er minal amino acid t o
it s cor r esponding t RNA. This is called t he p e p t i d yl -t RNA.
(b) The a mi n o a c yl s i t e or A-s i t e. This is t he sit e adjacent t o t he P-sit e
wher e t he next amino acyl-t RNA will bind at t he r ibosome. It is
posit ioned dir ect ly over t he t r iplet codon t o allow codon-ant icodon
pair ing.
(iii) At init iat ion, fMet -t RNA is in t he P-sit e and t he 2nd codon is posit ioned at
t he A-sit e.
2. El on ga t i on
(a) Thr ee r eact ions
(i) Bi n d i n g of aa-t RNA t o A-sit e
(a) Specificit y det er mined by codon, ant icodon pair ing.
(b) GTP is hydr olyzed at t he r ibosome is pair ing is cor r ect .
(c) GTP hydr olysis is par t of p r oofr e a d i n g.
(d) If ant i-codon and codon ar e complet ely non-complement ar y, amino
acyl-t RNA will not bind t o r ibosome.
(e) If complement is sufficient , binding and GTP hydr olysis occur .
(f) If complement is not per fect , t he r ibosome will r eject t he amino acyl-
t RNA.
(g) Hydr olysis of GTP must occur befor e t he next st ep in t r anslat ion can
occur .
(ii) Tr a n s p e p t i d a t i on
(a) Tr ansfer of t he gr owing polypept ide fr om pept idyl t RNA at t he P-sit e
t o t he amino acyl-t RNA at t he A-sit e.
(b) The est er bond bet ween t he pept ide and t RNA of t he pept idyl-t RNA is
br oken and t he car boxyl-t er minus of t he pept ide r eact s wit h t he amino
gr oup of t he amino acid of t he amino acyl-t RNA t o for m a pept ide bond
at t he A-sit e. The hydr olysis of t he est er bond pr ovides sufficient
ener gy t o dr ive pept ide bond for mat ion.
(c) Cat alyzed by p e p t i d yl t r a n s fe r a s e.
110 CSIR-NET Life Sciences
(iii) Tr a n s l oca t i on
(a) Movement of pept idyl-t RNA fr om A-sit e t o t he P-sit e and r elease of
d e a cyla t e d t RNA fr om t he P-sit e.
(b) Also r equir es GTP hydr olysis.
(iv) St eps in elongat ion ar e r epeat ed unt il a st op codon is r eached.
(v) Many r ibosomes at var ious st ages of t r anslat ion can be bound t o a single
mRNA.
(a) This st r uct ur e is called a polysome.
(b) As a r esult many polypept ides ar e synt hesized simult aneously fr om a
single mRNA.
3. Te r mi n a t i on
(a) When t he r ibosome encount er s a UGA, UAG or UAA codon, no amino acid is
added t o t he polypept ide. These codons ar e called t e r mi n a t i on cod on s .
(b) Ter minat ion also r equir es r e le a se fa ct or s.
(c) Funct ion of r elease fact or s
(i) Cat alyze hydr olysis of pept idyl t RNA
(ii) Pr omot e dissociat ion of 50S subunit
(d) 30S dissociat es or moves t o t he next st ar t codon on t he polycist r onic mRNA.
C. Eukaryotic translation
1. Sever al differ ences fr om pr okar yot es in t he init iat ion st age.
(a) I nit ia t ion t a kes pla ce a t 1st AUG on t he mRNA wit hin Koza k Sequence
(CCRCCAUGG).
(b) Met hionyl-t RNAmet is used t o init iat e t r anslat ion.
(c) Ther e is no Shine/Dalgar no sequence in eukar yot es. The 40S Rb binds t o t he 5'
cap st r uct ur e of t he mRNA and scans t o kozak sequence.
(d) Init iat ion complet e wit h associat ion of 60S subunit .
2. Elongat ion and t er minat ion ar e ver y similar in pr okar yot es and eukar yot es.
IV. Energetics of Translation
A. High ener gy bonds for synt hesis of polypept ide chain of (N) amino acid r esidues
N ATP t o char ge t RNA
1 GTP for init iat ion
N-1 GTP for binding of aa-t RNA t o A-sit e
N-1 GTP for t r anslocat ion st ep
1 GTP for t er minat ion
Sum: 3N
300 aa pr ot ein r equir es 900 high ener gy bonds; Each bond is 7.5 kcal/mol
900 bonds × 7.5 kcal/bond = 6,750 kcal/mol polypept ide
22.5 kcal/bond
Fr ee ener gy of pept ide bond hydr olysis is –0.5 kcal/mol
At 10% ener gy conver sion, –5 kcal would be sufficient t o for m a pept ide bond.
Biochemistry 111
B. High pr ice paid for synt hesizing a defined sequence.
1. Random polymer izat ion is cheaper
2. Only one high ener gy bond is involved in pept ide bond for mat ion
3. Cells need t o pr oofr ead t o insur e defined sequence.
4. The ult imat e st ep in gene expr ession must be cor r ect .
METABOLIC INTERRELATIONSHIPS
I. Review of digestion
A. Car bohydr at es, Pr ot eins hydr olyzed and cir culat ed as monomer s
1. N a
+
dependent upt ake int o int est inal cells
2. Intestinal cells ® blood ® liver ®
3. Insulin st imulat es upt ake
B. Fat digest ion differ ent fr om CHO, Pr ot ein
1. r equir es bile salt s; Vit . A, E, and K fat soluble
2. br eakdown, r esynt hesis of t r iglycer ides in mucosal cells
3. chylomicr ons ® LYMPH, by-passing liver
4. Medium Chain Fat t y Acids ent er blood dir ect ly as fr ee fat t y acids
5. Liver t akes up VLDL r emnant s, LDL, r esynt hesizes VLDL
6. Role of specific apolipopr ot eins
(a) A-I act ivat es HDL-LCAT (Lecit hin-Cholest . Acyl t r ansfer ase)
(b) B-48, B100, E for CM, VLDL, LDL r ecept or s
(c) C-II act ivat es Lipo Pr ot ein Lipase (LPL)
7. Effect s of int r acellular cholest er ol
(a) dir ect ly inhibit s and decr eases synt hesis r at e of HMG-CoA Reduct ase
(b) decr eases r at e of synt hesis of LDL r ecept or s
(c) st imulat es Acyl-CoA Cholest er ol Acyl Tr ansfer ase (ACAT)
II. Review of Metabolism, Regulation
A. Glycolysis, Gluconeogenesis
1. PFK-2 r egulat ion:
(a) cAMP-dependent PK inhibit s it in liver
(b) cAMP-dependent PK act ivat es it in hear t
(c) cAMP-dependent PK has no effect in skelet al muscle
2. Fut ile cycles and r egulat ion
B. Glycogenesis, Glycogenolysis: Ca
++
, cAMP-dependent phosphor ylat ions
C. Fat t y acid oxidat ion, fat t y acid synt hesis
1. cAMP ®® Hormone Sensitive Lipase, Acetyl CoA Carboxylase
2. Malonyl CoA blocks CAT transporter of FA into mitochondrion
D. Cit r ic Acid Cycle
1. NADH/NAD
+
r at io
112 CSIR-NET Life Sciences
2. Ca
++
act ivat es ICDH, a-KGDH, PDH
3. [Oxaloacet at e] oft en limit ing
E. Oxidat ive Phosphor ylat ion
1. ATP/ADP,Pi “Mass act ion r at io” cont r ols (DGATP)
2. ATP synt hesis in equilibr ium wit h Pr ot on Mot ive For ce (PMF)
F. Pent ose Phosphat e Pat hway
1. Impor t ant sour ce of NADPH, pent oses
2. [NADPH]/[NADP
+
] r at io cont r ols oxidat ive st eps
3. Pent ose-P / Hexose-P r at io cont r ols r ever sible st eps
G. Amino acid synt hesis, degr adat ion
1. Bot h r equir e aminot r ansfer ases, glut amic dehydr ogenase
2. Ala, Gln major amino acids for N t r anspor t in blood
3. Gln, Asp major Nit r ogen sour ces for biosynt hesis of ot her compounds (ur ea,
pur ines, pyr imidines, Asn, et c.)
III. Metabolic Division of Labor among Major Organs
A. Liver
1. Major pr oducer of fuel for ot her t issues: glucose, ket one bodies, VLDL
2. Uses glucose (fed), fat t y acids (fast ed), and amino acids as fuel (eit her )
(a) Essent ial amino acids cat abolized only when in excess (high Km enz.)
(b) BCAA ® muscle cat abolism
(c) gluconeogenesis fr om amino acids r equir es ur ea synt hesis
(d) gluconeogenesis dr iven by ener gy fr om fat t y acid oxidat ion
B. Adipose
1. The major t r iacylglycer ol (and calor ies) st or e
2. One of t he major fat es for excess diet ar y fat s
3. Synt hesis and br eakdown of t r iglycer ides under hor monal cont r ol
4. Synt hesis r equir es glucose (NADPH via PPP; glycer ol phosphat e)
5. Hor mone sensit ive lipase (cAMP) r eleases FA, glycer ol
C. Skeletal muscle
1. Fuel pr efer ence: Fat t y acids > ket one bodies > glucose; BCAA
2. Glucose and glycogen pr imar y fuel dur ing exer t ion
3. Ear ly st ar vat ion (fir st few days): muscle pr ot ein is br oken down for ener gy t hr oughout
body and for gluconeogenesis (liver )
4. Muscle r eleases amino gr oups as ALA, GLN
5. As ket one bodies fr om liver r ise, muscle pr ot eolysis shut off
D. Heart
1. Hear t has mor e mit ochondr ia per gm of t issue t han muscle, liver
Biochemistry 113
2. Fuel pr efer ence: Fat t y acids > ket one bodies > glucose
3. Negligible st or es of glycogen or t r iglycer ides: r equir es cont inual supply of fuels fr om
blood (glucose, fat t y acids, lact at e, ket one bodies)
E. Brain
1. Glucose pr efer r ed subst r at e (60% of t ot al glucose consumpt ion at r est )
2. Long t er m fast ing: adapt s t o ket one bodies as fuel (r at e depends on conc.)
F. Other tissues
1. Int est inal cells:
(a) Met abolize ASP, ASN, GLU, GLN for ener gy
(b) Release cit r ulline, pr oline, NH
3
, and ala
2. Kidney
(a) Cit r ulline fr om int est inal cells used t o make ARG
(b) Long t er m fast ing – significant gluconeogenesis
(c) In acidosis, GLN ® ALA + NH
4
+
(excr et ed) + HCO
3

(r et ained)
3. Red blood cells: GLUCOSE, GLUCOSE, GLUCOSE
Some Commonly Used Inhibitors of Protein Synthesis
In h i b i t or Su b u n i t St e p (s ) Re a ct i on Affe ct e d Cell Typ e
or F a c t or Bl ock e d Affect ed
Affect ed
Aur int r icar boxylic 30S/40S Init iat ion Binding of m-RNA Pr okar yot es/
acid Euka r yot es
Chlor a mphenicol 50 S Elongat ion Pept ide bond For mat ion Pr oka r yot es
Colcicin E3 30S Init iat ion Binding of m-RNA Pr oka r yot es
Elongat ion Binding of aminoacyl t -RNA
Cyclohexamide 60S Init iat ion Binding of Init iat or t -RNA Euka r yot es
Elongat ion Tr anslocat ion (t -RNA r elease
fr om P Sit e)
Dipt her ia Toxin eEF-2 Elongat ion Tr anslat ion Euka r yot es
Er t hr omycin 50 S Init iat ion For mat ion of Init iat ion complex Pr oka r yot es
Fusidic Acid EF-G/eEF-2 Elongat ion Tr anslocat ion Pr okar yot es/
Euka r yot es
Kasugamycin 30 S Init iat ion Binding of init iat or t -RNA Pr oka r yot es
Pur omycin 50S/60S Elongat ion Pept ide bond for mat ion (Tr igger Pr okar yot es/
immat ur e chain r elease) Euka r yot es
Spect inomycin 30 S Elongat ion Tr anslocat ion Pr oka r yot es
St r ept omycin 30 S Init iat ion Binding if Init iat or t -RNA Pr oka r yot es
Elongat ion Binding of aminoacyl t -RNA
(Induced Misr eading)
Tet r acycline 30 S Elongat ion Binding of aminoacyl t -RNA Pr oka r yot es
Ter minat ion Binding of RF-1 and RF-2
Practice Test Paper-I
1. The pH of t he blood is maint ained by
(a) miner al salt s (b) globulins
(c) albumins (d) haemoglobin
2. A pr ot ein having bot h st r uct ur al and enzymat ic t r ait s is
(a) act in (b) haemoglobin
(c) myosin (d) myoglobin
3. Pot assium cyanide st ops cell act ivit y by
(a) pr event ing diffusion of O
2
(b) blocking t r achea
(c) pr event ing t r ansfer of elect r ons fr om cyt ochr ome a3 t o O
2
(d) coagulat ing pr ot ein car r ier s
4. Successful Replicat ion of chr omosomes does not r equir e t he pr esence of
(a) Ribosomes (b) DNA Templat e
(c) Nuclear enzymes (d) ATP
5. In a diluted starch solution, a-salivar y amylase is added at pH 1.6 and kept at 35°C for
half an hour and t hen Iodine solut ion is added. What would be t he r esult ?
(a) no black colour will appear
(b) black colour will appear
(c) solut ion will be clear and colour less
(d) solut ion will be sweet
6. An amino acid devoid of a fr ee-COOH gr oup can be an
(a) Amine (b) Amide
(c) Imine (d) Pur ine
7. The most st r iking example of an unique point mut at ion is found in a disease called
(a) Night blindness (b) Thalassemia
(c) Down syndr ome (d) Sickle cell anaemia
8. For t he for mat ion of one molecule of hexose sugar , how many t ur ns of Calvin cycle
ar e needed ?
(a) Six (b) One
(c) Thir t y-six (d) One-sixt h
9. The hor mone t hat cont r ols pulse r at e and blood pr essur e is
(a) vasopr essin (b) oxyt ocin
(c) adr enalin (d) t hyr oxine
Practice Test Paper–Biochemistry 115
10. The maximum number of Hydr ogen bonds in which wat er molecule can par t icipat e
is?
(a) 1 (b) 2
(c) 3 (d) 4
11. Denat ur at ion of pr ot eins leads t o loss of biological act ivit y by
(a) for mat ion of amino acid
(b) loss of pr imar y st r uct ur e
(c) loss of bot h pr imar y and secondar y st r uct ur e
(d) loss of secondar y and t er t iar y st r uct ur e
12. The pKa of acet ic acid is 4.76. What is t he pH of 0.1 M solut ion of acet ic acid?
(a) 5.76 (b) 2.88
(c) 2.43 (d) 1.88
13. 40 g of ice at 0 °C and 80 g wat er at 40
o
C ar e mixed t hor oughly, t he t emper at ur e of
t he mixt ur e will be
(a) 0° C (b) 10 °C
(c) 20 °C (d) 30 °C
14. Galact ose and Mannose ar e t wo isomer ic monosacchar ides, which ar e
(a) Enant iomer s (b) Epimer s
(c) Anomer s (d) Diast er eoisor ner s
15. Pr ot eins absor bs maximally UV r adiat ion at 280 nm due t o t he pr esence of
(a) Tr ypt ophan (b) Tyr osine
(c) Phenyl alanine (d) All of t he above
16. Which one of t he following solut ions will have t he least Osmot ic Pr essur e?
(a) 0.5 M glucose (b) 0.5 M NaCl
(c) 0.5 M Na
2
SO
4
(d) 0.499 M St ar ch
17. The fr ee ener gy cha nge (DG)at t he midpoint of helix-coil t r ansit ion in DNA melt ing
cur ve is equal t o
(a) zer o (b) 1.0
(c) –1.0 (d) indet er minat e
18. DNA is usually dissolved in TRIS-EDTA buffer , EDTA is added t o ensur e t hat
(a) DNA is not degr aded by nucleases
(b) Divalent met al ions ar e chelat ed
(c) Bot h of t he above
(d) pH is kept alkaline
19. Two pr ot eins have same molecular weight and same isoelect r ic point . The best way
t o r esolve t hem will be by
(a) Ion Exchange Chr omat ogr aphy
(b) Gel Filt r at ion Chr omat ogr aphy
(c) Rever se Phase Chr omat ogr aphy
(d) Chr omat ofocussing
116 CSIR-NET Life Sciences
20. Mar k t he incor r ect st at ement : Ribozyme and Abzyme ar e
(a) bot h enzymes
(b) bot h pr ot eins
(c) RNA and pr ot ein r espect ively
(d) able t o hydr olyse phosphodiest er and pept ide bonds r espect ively
21. DNA sequencing is done on a sequencing gel, which is
(a) SDS-PAGE (b) Ur ea-PAGE
(c) Nat ive PAGE (d) Agar ose
22. The digest ion of lipid t akes place in small int est ine by lipases, However , it does not
occur t ill bile fluid mixes wit h t he food. This mixing up of food wit h bile juice is
essent ial for digest ion of lipids because
(a) lipases pr esent in t he int est ine r equir e anot her coenzyme pr esent in t he bile
juice for it s funct ioning
(b) Emulsificat ion is must for fat s t o be digest ed and for lipases t o act upon lipids
(c) lipase can only act at alkaline pH
(d) None of t he above
23. Mar k t he Incor r ect st at ement :
(a) Endocr ine signaling is r elat ively slow because it depends on diffusion and blood
flow
(b) If t he same hor mone has differ ent effect s on differ ent t ar get or gans, t he r ecept or
ar e usually differ ent
(c) Wat er soluble hor mones int er act wit h cell sur face r ecept or s, wher eas lipid soluble
hor mones usually bind t o int r acellular r ecept or s
(d) The specificit y of hor mone signaling depends on r esponsive cells possessing
r ecept or s t hat bond t he hor mone t ight ly by means of mult iple weak bonds.
24. Silencer s ar e locat ed t o upst r eam of t r anscr ipt ion st ar t sit e and funct ion-
(a) in bot h t he or ient at ion (b) in for war d or ient at ion
(c) in r ever se or ient at ion (d) None of t he above
25. The r enat ur at ion of t he genome is a r andom collision of complement ar y sequences
and follows
(a) fir st or der kinet ics (b) second or der kinet ics
(c) t hir d or der kinet ics (d) None of t he above
26. Which one of t he following elect r ophor esis dependent s least on t he char ge of t he
pr ot ein?
(a) Fr ee zone capillar y elect r ophor esis
(b) Gel elect r ophor esis
(c) SDS-polycr ylamide gel elect r ophor esis
(d) Isoelect r ic focussing
Practice Test Paper–Biochemistry 117
27. Which one of t he following compound has t he highest gr oup t r ansfer pot ent ial for
phosphat e ?
(a) Glucose-6-phosphat e (b) Fr uct ose-1, 6-diphosphat e
(c) 2-phosphoenolpyr uvat e (d) Adenosine t r iphosphat e
28. Which one of t he following set s of glycolyt ic enzymes is allost er ically r egulat ed?
(a) Glucokinase, phosphofr uct okinase and pyr uvat e kinase
(b) Hexokinase, aldolase and pyr uvat e kinase
(c) Phosphofr uct okinase, enolase and pyr uvat e kinase
(d) Hexokinase, phosphofr uct okinase and pyr uvat e kinase
29. Which one of t he following compounds is r equir ed for acet yl coenzyme A (acet yl
CoA)t o ent er t he cit r ic acid cycle?
(a) Isocit r at e (b) Malat e
(c) Oxaloacet at e (d) Pyr uvat e
30. Which one of t he following DNA polymer ase is essent ial for bot h t he r eplicat ion and
r epair of DNA?
(a) DNA polymer ase I (b) DNA polymer ase II
(c) DNA polymer ase III (d) DNA polymer ase d
31. The fact or r equir ed only for accur at e init iat ions of t r anscr ipt ion in pr okar yot es is
(a) alpha (a) (b) bet a (b)
(c) r ho (r) (d) sigma (s)
32. A post -t r anscr ipt ional pr ocessing event t hat occur s in t he for mat ion of bot h messenger
RNA (mRNA)and t r ansfer RNA (t -RNA) in eukar yot es is
(a) t he addit ion of t he sequence CCA t o 3' ends
(b) t he r emoval of int r ons by splicing
(c) t he modificat ion of some of t heir bases
(d) cleavage event befor e polyadenylat ion
33. Eukar yot ic mRNAs differ fr om Pr okar yot ic mRNAs in t hat
(a) t hey do not have a 5' unt r anslat ed r egion
(b) t heir coding r egions ar e separ at ed by spacer s
(c) t hey do not have 3' UTR
(d) t hey have a fr ee 3' hydr oxy gr oup on each of t heir ends
34. Degener acy of t he genet ic code means t hat
(a) a given base t r iplet can code for mor e t han one amino acid
(b) t her e is no punct uat ion in t he code sequence
(c) t he t hir d base in a codon is net impor t ant for coding
(d) a given amino acid can be coded for by mor e t han one base t r iplet
35. Which one of t he following sequences is most likely t o be a r est r ict ion enzyme
r ecognit ion sit e?
(a) CGGC (b) CGC
(c) GTAATG (d) GTCGAC
118 CSIR-NET Life Sciences
36. Which st ep is t he fir st in t he for mat ion of a double st r anded complement ar y DNA
(c DNA) t ar get for cloning?
(a) blunt end ligat ion
(b) DNA dependent DNA synt hesis
(c) Rest r ict ion enzyme cleavage
(d) Pr imer annealing
37. Cyclic AMP r egulat es t he lact ose (lac)oper on by
(a) binding t o t he oper at or t o t ur n on t r anscr ipt ion
(b) binding t o t he lac r epr essor t o pr event t r anscr ipt ion
(c) combining wit h t he cat abolit ic act ivat or pr ot ein (CAP) t o for m a complex t hat
enhances t r anscr ipt ions upon binding t o t he pr omot or .
(d) combining wit h t he CAP t o r emove CAP’s inhibit ion of t r anscr ipt ion
38. A 4 Kb covalent ly closed cir cular (ccc) plasmid has 2 (t wo) sit es for a r est r ict ion enzyme
1 kb apar t . It is par t ially digest ed by t he enzyme. The maximum number of bands
t hat could be seen on a gel will be
(a) 1 (b) 2
(c) 3 (d) 4
39. In an enzyme r eact ion t he r eact ion velocit y becomes mor e t han double when t he
subst r at e concent r at ion is doubled. This is possible when t he equat ion gover ning t he
kinet ics is
(a) Michelis-Ment on Kinet ics
(b) Micheils-Ment on Kinet ics wit h subst r at e inhibit ion
(c) Michelis-Ment on Kinet ics wit h pr oduct inhibit ion
(d) Hill Equat ion
40. A mixt ur e separ at ed on HPLC gives clear ly defined peaks wher eas in a manually r un
column t he peaks t end t o blend int o each ot her . This is pr imar ily because
(a) t he small size of t he packing mat er ial used in t he HPLC column
(b) t he bet t er cont r ol of flow in HPLC
(c) t he use of high pr essur e in HPLC
(d) t he use of bet t er det ect ion syst ems in HPLC
41. Which is t r ue about enzymes,
(a) t hey always incr ease r at e of r eact ion
(b) t hey always decr ease r at e of r eact ion
(c) t hey do not dist ur b t he equilibr ium
(d) Always car r y ir r ever sible r eact ions
42. Linkage pr esent in cellulose molecule is
(a) b (1 ® 4) (b) a ( 1 ® 4)
(c) a (1 ®6) (d) bot h (b)and (c)
Practice Test Paper–Biochemistry 119
43. Blocking act ion of enzyme t hr ough blocking it s act ive sit e is
(a) Allost er ic inhibit ion (b) Feedback inhibit ion
(c) Compet it ive inhibit ion (d) Non-compet it ive inhibit ion
44. Which one of t he following is wit hout coenzyme act ivit y ?
(a) Vit amin E (b) Thiamine
(c) Biot in (d) Riboflavin
45. Act ive t r anspor t
(a) Releases ener gy (b) Requir e ener gy
(c) Pr oduces ener gy (d) Pr oduces t oxic mat er ial
46. A shor t lengt h of double st r anded DNA molecule cont ains 120 adenine and 120 cyt osine
bases. The t ot al number of nucleot ides in t his DNA fr agment is
(a) 60 (b) 120
(c) 240 (d) 480
47. Which of t he following element plays an impor t ant r ole in nit r ogen fixat ion?
(a) Manganese (b) Molybdenum
(c) Zinc (d) Copper
48. Specificit y of an enzyme depends upon
(a) Act ive sit e (b) Linear sequence
(c) Km (d) Tur n over number
49. Sodium Dodecyl Sulphat e (SDS) is used while separ at ing pr ot eins by polyacr ylanide
gel elect r ophor esis because
(a) It helps in solubilizat ion of pr ot eins t her eby making it easier t o separ at e
(b) It binds t o pr ot eins and confer s unifor m negat ive char ge densit y t her eby making
t hem move dur ing elect r ophor esis
(c) Decr eases t he sur face t ension of t he buffer used for elect r ophor esis
(d) St abilizes t he pr ot eins
50. In a subst it ut ion nucleophilic fir st or der r eact ion (SN1)t he st er eochemical out come
is
(a) Raemisat ion (b) Inver sion of configur at ion
(c) Ret ent ion of configur at ion (d) Difficult t o pr edict
51. Which of t he following chemot her a peut ic a gent s wor ks by impa ir ing de novo
pur inesynt hesis?
(a) acyclovir (acycloguanosine) (b) 5-fluor our acil (ant imet abolit e)
(c) met hot r exat e (ant i folat e) (d) hydr oxyur ea
52. The lar gest ener gy r eser ve (in t er ms of kilocalor ies) in humans is:
(a) liver glycogen (b) muscle glycogen
(c) adipose t issue t r iacylglycer ol. (d) muscle pr ot ein.
120 CSIR-NET Life Sciences
53. In t he int er act ion of a hor mone wit h it s r ecept or all of t he following ar e t r ue EXCEPT:
(a) mor e t han one polypept ide chain of t he hor mone may be necessar y.
(b) mor e t han one second messenger may be gener at ed.
(c) an ar r ay of t r ansmembr ane helices may for m t he binding sit e for t he hor mone.
(d) hor mones r eleased fr om t heir r ecept or aft er endocyt osis could t heor et ically
int er act wit h a nuclear r ecept or .
54. Some hor mone-r ecept or complexes ar e int er nalized by endocyt osis. This pr ocess may
involve:
(a) binding of hor mone-r ecept or complex t o a clat hr in coat ed pit .
(b) r ecycling of r ecept or t o cell sur face.
(c) for mat ion of a r ecept osome.
(d) all of t he above
55. In hypopit uit ar ism it is necessar y t o maint ain t he ovar ian cycle in female pat ient s. In
t he ovar ian cycle:
(a) GnRH ent er s t he vascular syst em via t r anspor t by a specific membr ane car r ier .
(b) cor pus lut eum r apidly involut es only if fer t ilizat ion does not occur .
(c) inhibin wor ks by inhibit ing synt hesis of a subunit of FSH.
(d) LH is t aken up by cor pus lut eum and binds t o cyt oplasmic r ecept or s.
56. All of t he following st at ement s about act in and myosin ar e t r ue EXCEPT:
(a) t he globular head sect ion of myosin has domains for binding ATP and act in.
(b) act in is t he major pr ot ein of t he t hick filament .
(c) t he binding of ATP t o t he act in-myosin complex pr omot es dissociat ion of act in
and myosin.
(d) F-act in, for med by aggr egat ion of G-act in-ATP-MgH complex, is st abilized when
t r opomyosin is bound t o it .
57. St ar ch digest ion is mor e efficient aft er heat ing t he st ar ch wit h wat er because heat ing:
(a) hydr at es t he st ar ch gr anules, making t hem mor e suscept ible t o pancr eat ic
amylase.
(b) par t ly hydr olyses a-1, 6 links.
(c) conver t s t he linear amylose t o br anched amylopect in, which r esembles glycogen.
(d) inact ivat es amylase inhibit or s, which ar e common in t he t issues of st ar chy plant s.
58. Micelles:
(a) ar e t he same as emulsion dr oplet s.
(b) for m fr om bile acids at all bile acid concent r at ions.
(c) alt hough t hey ar e for med dur ing lipid digest ion, do not significant ly enhance
ut ilizat ion of diet ar y lipid.
(d) always consist of only a single lipid species.
Practice Test Paper–Biochemistry 121
59. Cer t ain t issues effect Cl

secr et ion via a Cl

channel (CFTR pr ot ein-cyst ic fibr osis
t r ansmembr ane r egulat or y pr ot ein). Choler a t oxin abnor mally opens t he channel
leading t o a loss of NaCl. Tr eat ment for choler a and spor t s dr inks for elect r olyt e
r eplacement ar e fluids high in Na
+
and glucose. The pr esence of glucose enhances
NaCl r eplenishment because:
(a) absor bing any nut r ient causes Na
+
upt ake.
(b) glucose pr event s Na
+
excr et ion.
(c) Na
+
and glucose ar e t r anspor t ed in opposit e dir ect ions.
(d) glucose is a bsor bed a cr oss int est ina l epit helia l cells via a Na
+
-dependent
cot r anspor t er .
60. Of t wo people wit h appr oximat ely t he same weight , t he one wit h t he higher basal
ener gy r equir ement would most likely be:
(a) t aller (b) female if t he ot her wer e male.
(c) older (d) under less st r ess.
61. Basal met abolic r at e:
(a) is not influenced by ener gy int ake.
(b) may decr ease up t o 50% dur ing per iods of st ar vat ion.
(c) incr eases in dir ect pr opor t ion t o daily ener gy expendit ur e.
(d) is not r esponsive t o changes in hor mone levels.
62. The pr imar y effect of t he consumpt ion of excess pr ot ein beyond t he body’s immediat e
needs will be:
(a) excr et ion of t he excess as pr ot ein in t he ur ine.
(b) an incr ease in t he “st or age pool” of pr ot ein.
(c) an incr eased synt hesis of muscle pr ot ein.
(d) an enhancement in t he amount of cir culat ing plasma pr ot eins.
63. Kwashior kor is:
(a) t he most common for m of pr ot ein-calor ie malnut r it ion in t he INDIA.
(b) char act er ized by a t hin, wast ed appear ance.
(c) an adequat e int ake of t ot al calor ies but a specific deficiency of pr ot ein.
(d) an adequat e int ake of t ot al pr ot ein but a deficiency of t he essent ial amino acids.
64. A complet e r eplacement of animal pr ot ein in t he diet by veget able pr ot ein:
(a) would be expect ed t o have no effect at all on t he over all diet .
(b) would r educe t he t ot al amount of food consumed for t he same number of calor ies.
(c) might r educe t he t ot al amount of ir on and vit amin B12 available.
(d) would be sat isfact or y r egar dless of t he nat ur e of t he veget able pr ot ein used.
65. Diet ar y fat :
(a) is usually pr esent , alt hough t her e is no specific need for it .
(b) if pr es en t in exces s , ca n be s t or ed a s eit h er glycogen or a dipos e t is s u e
t r iacylglycer ol.
(c) should include linoleic and linolenic acids.
(d) should incr ease on an endur ance t r aining pr ogr am in or der t o incr ease t he body’s
ener gy st or es.
122 CSIR-NET Life Sciences
66. For diabet ics:
(a) t he only car bohydr at e t hat must be eliminat ed in t he diet is sucr ose.
(b) not all car bohydr at e foods r aise blood glucose levels at t he same r at e because
t he glycemic index of all foods is not t he same.
(c) who ar e nor mally in good cont r ol, st r ess will have no effect on t heir blood sugar
levels.
(d) a veget ar ian diet is t he only appr opr iat e choice.
67. The effect s of vit amin A may include all of t he following EXCEPT:
(a) pr event ion of anemia. (b) ser ving as an ant ioxidant .
(c) cell differ ent iat ion. (d) t he visual cycle.
68. Ascor bic acid may be associat ed wit h all of t he following EXCEPT:
(a) ir on absor pt ion.
(b) bone for mat ion.
(c) wound healing.
(d) par t icipat ion in hydr oxylat ion r eact ions.
69. The t r ansmembr ane por t ion of a pr ot ein spanning t he membr ane is most likely t o be
composed of aminoacids t hat ar e-
(a) basic (b) acidic
(c) glycosylat ed (d) hydr ophobic
70. All t he following compounds ar e Int er mediat e of t he cit r ic acid cycle except
(a) Isocit r at e (b) Malat e
(c) Pyr uvat e (d) Succinat e
71. Cellulose is a _____ ma de of ma ny _____.
(a) polypept ide . . . monomer s
(b) car bohydr at e . . . fat t y acids
(c) polymer . . . glucose molecules
(d) pr ot ein . . . amino acids
72. The four main cat egor ies of macr omolecules in a cell ar e
(a) pr ot eins, DNA, RNA, and st er oids.
(b) monosacchar ides, lipids, polysacchar ides, and pr ot eins.
(c) pr ot eins, nucleic acids, car bohydr at es, and lipids.
(d) nucleic acids, car bohydr at es, monosacchar ides, and pr ot eins.
73. The char act er ist ic t hat all lipids have in common is
(a) t hey ar e all made of fat t y acids and glycer ol.
(b) none of t hem is ver y high in ener gy cont ent .
(c) t hey ar e all acidic when mixed wit h wat er .
(d) none of t hem dissolves in wat er .
Practice Test Paper–Biochemistry 123
74. How you cellulose and st ar ch ar e differ ent
(a) Cellulose molecules ar e much t oo lar ge.
(b) St ar ch is made of glucose; cellulose is made of ot her sugar s.
(c) The bonds bet ween sugar s in cellulose ar e much st r onger .
(d) The sugar s in cellulose bond t oget her differ ent ly t han in st ar ch, giving cellulose
a differ ent shape.
75. In some places a pr ot ein molecule may t wist or fold back on it self. This is called
_____ and t he coils or folds ar e held in place by _____.
(a) t er t iar y st r uct ur e . . . hydr ogen bonds
(b) pr imar y st r uct ur e . . . covalent bonds
(c) secondar y st r uct ur e . . . pept ide bonds
(d) t er t iar y st r uct ur e . . . covalent bonds
76. A hydr ophobic amino acid R gr oup would be found wher e in a pr ot ein?
(a) for ming a pept ide bond wit h t he next amino acid in t he chain
(b) on t he out side of t he folded chain, in t he wat er
(c) on t he inside of t he folded chain, away fr om wat er
(d) for ming hydr ogen bonds wit h ot her R gr oup
77. The over all t hr ee-dimensional shape of a polypept ide is called t he
(a) double helix. (b) pr imar y st r uct ur e.
(c) secondar y st r uct ur e. (d) t er t iar y st r uct ur e.
78. How many differ ent kinds of pr ot ein molecules ar e t her e in a t ypical cell?
(a) Twent y (b) about a hundr ed
(c) Thousands (d) billions
79. The building blocks of nucleic acid molecules ar e called
(a) polysacchar ides. (b) amino acids.
(c) fat t y acids. (d) nucleot ides.
80. Which of t he following do nucleic acids and pr ot eins have in common?
(a) They ar e bot h made of amino acids.
(b) Their st r uct ur es cont ain sugar s.
(c) They ar e hydr ophobic.
(d) They ar e lar ge polymer s.
81. Which of t he following would pr obably not be affect ed when a pr ot ein is denat ur ed?
(a) pr imar y st r uct ur e (b) secondar y st r uct ur e
(c) hydr ogen bonds (d) t er t iar y st r uct ur e
82. A glucose molecule is t o st ar ch as
(a) a st er oid is t o a lipid. (b) a pr ot ein is t o an amino acid.
(c) a nucleic acid is t o a polypept ide. (d) a nucleot ide is t o a nucleic acid.
124 CSIR-NET Life Sciences
83. Palm oil and coconut oil ar e mor e like animal fat s t han ot her plant oils. Because t hey
____ t han ot her plant oils, t hey can cont r ibut e t o car diovascular disease.
(a) cont ain fewer double bonds (b) ar e less sat ur at ed
(c) cont ain mor e sodium (d) ar e less soluble in wat er
84. A shor t age of phosphor us in t he soil would make it especially difficult for a plant t o
manufact ur e
(a) DNA (b) pr ot eins.
(c) cellulose. (d) fat t y acids.
85. How does DNA differ fr om RNA?
(a) DNA is lar ger .
(b) One of t heir nit r ogenous bases is differ ent .
(c) They cont ain differ ent sugar s.
(d) DNA consist s of t wo st r ands in a double helix.
86. Which of t he following r anks t he molecules in t he cor r ect or der by size?
(a) wat er . . . sucr ose . . . glucose . . . pr ot ein
(b) pr ot ein . . . wat er . . . glucose . . . sucr ose
(c) wat er . . . pr ot ein . . . sucr ose . . . glucose
(d) pr ot ein . . . sucr ose . . . glucose . . . wat er
87. On t he basis of t he pr inciple of complement ar y base pair ing you would expect t he
per cent age of ___ t o be equal t he per cent age of ____.
(a) A ... T (b) A ... G
(c) T ... G (d) A ... C
88. Lipids differ fr om ot her lar ge biological molecules in t hat t hey
(a) ar e much lar ger . (b) ar e not t r uly polymer s.
(c) do not have specific shapes. (d) do not cont ain car bon.
89. Wat er is a polar molecule. This means t hat
(a) t he opposit e ends of t he molecule have opposit e char ges.
(b) wat er molecules ar e linear , like a pole.
(c) wat er is one of t he many hydr ophobic molecules.
(d) t he at oms in wat er have elect r onegat ivit es which ar e equal.
90. Cellular r espir at ion is an example of
(a) an anabolic pat hway. (b) a cat abolic pat hway.
(c) bioener get ics. (d) t her modynamics.
91. Or ganisms ar e descr ibed as t her modynamically open syst ems. This means t hat
(a) t he met abolism of an or ganism is isolat ed fr om it s sur r oundings
(b) or ganisms can r ever se t he incr ease in ent r opy.
(c) or ganisms acquir e ener gy fr om t heir sur r oundings.
(d) or ganisms ar e capable of cir cumvent ing t he second law of t her modynamics.
Practice Test Paper–Biochemistry 125
92. Fr om t he equat ion delt a G = delt a H -T delt a S it is clear t hat
(a) a decr ease in t he syst em’s t ot al ener gy will incr ease t he pr obabilit y of spont aneous
change.
(b) spont aneous change is most pr obable when t he syst em exper iences an incr ease
in ent r opy.
(c) incr easing t he t emper at ur e of a syst em will incr ease t he pr obabilit y of spont aneous
change.
(d) t he capacit y of a syst em t o per for m wor k is r elat ed t o t he t ot al ener gy of t he
syst em.
93. An ender gonic r eact ion is char act er ized by
(a) it s t endency t o occur spont aneously.
(b) t he net r elease of fr ee ener gy.
(c) a value for delt a G which is posit ive.
(d) r eact ant s t hat ar e mor e complex t han pr oduct s.
94. Cells cont inue t o funct ion only when a met abolic disequilibr ium is in effect ; cells
avoid r eaching met abolic equilibr ium
(a) because cellular met abolism ut ilizes only t hose r eact ions t hat ar e nonr ever sible.
(b) because t he pr oduct s of one r eact ion become t he r eact ant s for a differ ent r eact ion
and ar e unable t o accumulat e.
(c) because t he major it y of impor t ant met abolic r eact ions have delt a Gs of zer o.
(d) by limit ing t he ent r y of high fr ee ener gy molecules once t hey ha ve been
cat abolized.
95. What must be t r ue if t he r eact ion AB plus CD t o AC plus BD occur s spont aneously?
(a) The delt a G of t he r eact ion must be negat ive.
(b) t he r eact ion must be exer gonic.
(c) t he envir onment s has adequat e t her mal ener gy t o meet t he EA r equir ement .
(d) all of t he above
96. What best char act er izes t he r ole of ATP in cellular met abolism?
(a) The r elease of fr ee ener gy dur ing t he hydr olysis of ATP heat s t he sur r ounding
envir onment .
(b) I t s fr ee ener gy is coupled t o a n ender gonic pr ocess via t he for ma t ion of a
phosphor ylat ed int er mediat e.
(c) It is cat abolized t o car bon dioxide and wat er .
(d) The delt a G associat ed wit h it s hydr olysis is posit ive.
97. Enzymes ar e descr ibed as cat alyst s, which means t hat t hey
(a) pr ovide act ivat ion ener gy for t he r eact ions t hey facilit at e.
(b) change t he r at e of a r eact ion wit hout being consumed by t he r eact ion
(c) st abilize molecules in t he t r ansit ion st at e.
(d) elevat e t he EA bar r ier so t he molecules will not spont aneously degr ade.
126 CSIR-NET Life Sciences
98. What best explains t he obser vat ion of subst r at e specificit y?
(a) Ther e is a pr ecise compat ibilit y bet ween an enzyme’s act ive sit e and t he subst r at e
molecule
(b) Molecules and act ive sit es var y in size; only pr oper ly sized molecules can fit .
(c) React ion-specific enzymes, such as hydr olases, assume a fit by folding ar ound
t he most numer ous subst r at e molecules.
(d) Pola r it y compa t ibilit ies; a ct ive sit es cont a in elect r onega t ive a t oms while
subst r at es t end t o car r y slight posit ive char ges.
99. Sucr ase nor mally cat alyses t he cat abolism of sucr ose t o glucose and fr uct ose. What ,
if any, changes could be made t o t he syst em so t hat sucr ase could synt hesize sucr ose
fr om t he t wo pr oduct s?
(a) incr ease t he t emper at ur e of t he syst em
(b) alt er t he concent r at ion of molecules so t hat sucr ase is low and glucose and
fr uct ose ar e ver y high
(c) select ively denat ur e t he enzyme by var ying t he pH
(d) phosphor ylat e eit her t he glucose or t he fr uct ose
100. A plot of enzyme velocit y against t emper at ur e for an enzyme indicat es lit t le act ivit y
at 0 degr ees celsius and 45 degr ees celsius, wit h peak act ivit y at 35 degr ees celsius.
The most r easonable explanat ion for t he low velocit y at 0 degr ees celsius is t hat at
t his t emper at ur e
(a) t he hydr ogen bonds t hat define t he enzyme’s act ive sit e ar e unst able.
(b) at low t emper at ur es t he subst r at e becomes an allost er ic r egulat or .
(c) t he enzyme was denat ur ed.
(d) t her e is t oo lit t le act ivat ion ener gy available.
Practice Test Paper-II
1. Pr okar yot ic cells, but not eukar yot ic cells, have:
(a) endoplasmic r et iculum. (b) hist ones.
(c) nucleoid. (d) a nucleus.
2. Fact or s r esponsible for a wat er molecule being a dipole include:
(a) t he t et r ahedr al st r uct ur e of liquid wat er .
(b) t he magnit ude of t he H-O-H bond angle.
(c) t he abilit y of wat er t o hydr ogen bond t o var ious chemical st r uct ur es.
(d) t he differ ence in bond st r engt h bet ween hydr ogen bonds and covalent bonds.
3. Hydr ogen bonds can be expect ed t o for m only bet ween elect r onegat ive at oms such as
oxygen or nit r ogen and a hydr ogen at om bonded t o:
(a) car bon. (b) an elect r onegat ive at om.
(c) hydr ogen. (d) iodine.
4. The ion pr oduct of wat er :
(a) is independent of t emper at ur e.
(b) has a numer ical value of 1 × 10
–14
at 25°C.
(c) is t he equilibr ium const ant for t he r eact ion H
3
O
+
and OH

.
(d) r equir es t hat [H
+
] and [OH

] always be ident ical.
5. Lysosomal enzymes:
(a) ar e hydr olases.
(b) usually oper at e at acidic pH.
(c) ar e nor mally isolat ed fr om t heir subst r at es by t he imper meable lysosomal
membr a ne.
(d) All of t he above ar e cor r ect .
6. In a DNA double helix:
(a) t he individual st r ands ar e not helical.
(b) hydr ogen bonds for m bet ween a pur ine and a pyr imidine base on t he same st r and.
(c) adenine on one st r and is hydr ogen bonded t o t hymine on t he opposit e st r and.
(d) phospho diest er bonds ar e or ient ed t owar d t he int er ior of t he helix.
7. The A helix of DNA differ s fr om t he B helix in all of t he following EXCEPT:
(a) appear ance of t he major and minor gr ooves.
(b) pit ch of t he base pair s r elat ive t o t he helix axis.
(c) t hickness of t he helix.
(d) polar it y of t he st r ands.
8. The Z-DNA helix:
(a) has fewer base pair s per t ur n t han t he B-DNA.
(b) is favor ed by an alt er nat ing GC sequence.
128 CSIR-NET Life Sciences
(c) t ends t o be found at t he 3'-end of genes.
(d) is inhibit ed by met hylat ion of t he bases.
9. RNA:
(a) incor por at es bot h modified and unmodified pur ine and pyr imidine bases dur ing
t r anscr ipt ion.
(b) does not exhibit any double-helical st r uct ur e.
(c) st r uct ur es exhibit base st acking and hydr ogen-bonded base pair ing.
(d) usually cont ains about 65-100 nucleot ides.
10. Bent DNA:
(a) occur s only in t he pr esence of ext er nal agent s like t he ant it umor dr ugs.
(b) may be a fundament al element in t he int er act ion bet ween DNA sequences and
pr ot eins for such pr ocesses as t r anscr ipt ion and sit e-specific r ecombinat ion.
(c) occur s pr imar ily in t he pr esence of t r iple-st r anded DNA.
(d) r equir es t he pr esence of inver t ed r epeat s.
11. Chaper one pr ot eins:
(a) all r equir e ATP t o exer t t heir effect .
(b) cleave incor r ect disulfide bonds, allowing cor r ect ones t o subsequent ly for m.
(c) guide t he folding of polypept ide chains int o pat t er ns t hat would be t her mo-
dynamically unst able wit hout t he pr esence of chaper ones.
(d) of t he hsp70 class ar e involved in t r anspor t of pr ot eins acr oss mit ochondr ial and
endoplasmic r et iculum membr anes.
12. Pr ot eins may be separ at ed accor ding t o size by:
(a) isoelect r ic focusing.
(b) polyacr ylamide gel elect r ophor esis.
(c) ion-exchange chr omat ogr aphy
(d) molecular exclusion chr omat ogr aphy.
13. Changes in pr ot ein confor mat ion can be det ect ed r apidly by:
(a) ult r aviolet absor bance spect r oscopy.
(b) fluor escence emission spect r oscopy
(c) opt ical r ot at or y disper sion.
(d) all of above.
14. Which of t he following st at ement s about E. coli DNA polymer ases is cor r ect ?
(a) All polymer ases have bot h 3' t o 5' and 5' t o 3' exonuclease act ivit y.
(b) The pr imar y r ole of polymer ase III is in DNA r epair .
(c) Polymer ases I and III r equir e bot h a pr imer and a t emplat e.
(d) Polymer ase I t ends t o r emain bound t o t he t emplat e unt il a lar ge number of
nucleot ides have been added.
15. Bot h st r ands of DNA ser ve as t emplat es concur r ent ly in:
(a) r eplicat ion. (b) excision r epair .
(c) mismat ch r epair . (d) t r anscr ipt ion-coupled r epair .
Practice Test Paper–Biochemistry 129
16. All of t he following st at ement s about t elomer ase ar e cor r ect EXCEPT:
(a) t he RNA component act s as a t emplat e for t he synt hesis of a segment of DNA.
(b) it adds t elomer es t o t he 5' ends of t he DNA st r ands.
(c) it pr ovides a mechanism for r eplicat ing t he ends of linear chr omosomes in most
euka r yot es.
(d) it is a r ever se t r anscr ipt ase.
17. A t r ansit ion mut at ion:
(a) occur s when a pur ine is subst it ut ed for a pyr imidine, or vice ver sa.
(b) r esult s fr om t he inser t ion of one or t wo bases or base analogs int o t he DNA
chain.
(c) decr eases in fr equency in t he pr esence of base analogs such as 2-amino pur ine.
(d) r esult s fr om t he subst it ut ion of one pur ine for anot her or of one pyr imidine for
anot her .
18. Homologous r ecombina t ion:
(a) occur s only bet ween t wo segment s fr om t he same DNA molecule.
(b) r equir es t hat a specific DNA sequence be pr esent .
(c) r equir es t hat one of t he duplexes under going r ecombinat ion be nicked in bot h
st r ands.
(d) may r esult in st r and exchange by br anch migr at ion.
19. Of t he following ar e t r ue about t r ansposit ions EXCEPT:
(a) t r ansposons move fr om one locat ion t o a differ ent one wit hin a chr omosome.
(b) bot h t he donor and t ar get sit es must be homologous.
(c) t r ansposons have inser t ion sequences t hat ar e r ecognized by t r ansposases.
(d) t he t r ansposon may eit her be excised and moved or be r eplicat ed wit h t he
r eplicat ed piece moving.
20. All of t he following ar e t r ue about nucleot ide excision r epair EXCEPT:
(a) r emoval of t he damaged bases occur s on only one st r and of t he DNA.
(b) it r emoves t hymine dimer s gener at ed by UV light .
(c) it involves t he act ivit y of an excision nuclease, which is an endonuclease.
(d) only t he damaged nucleot ides ar e r emoved.
21. Dur ing t he elongat ion st age of eukar yot ic pr ot ein synt hesis:
(a) t he incoming aminoacyl-t RNA binds t o t he P sit e.
(b) a new pept ide bond is synt hesized by pept idyl t r ansfer ase sit e of t he lar ge
r ibosomal subunit in a GTP-r equir ing r eact ion.
(c) t he pept ide, st ill bound t o a t RNA molecule, is t r anslocat ed t o a differ ent sit e on
t he r ibosome.
(d) st r ept omycin can cause pr emat ur e r elease of t he incomplet e pept ide.
22. Ter minat ion of pr ot ein synt hesis:
(a) r equir es a st op codon t o be locat ed at t he P sit e of t he lar ge r ibosomal subunit .
(b) occur s when a non-r ibosomal pr ot ein r elease fact or binds t o t he r ibosome.
130 CSIR-NET Life Sciences
(c) r equir es t he act ion of a non-r ibosomal hydr olase t o r elease t he pept ide.
(d) does not r equir e ener gy.
23. 4-Hydr oxylat ion of specific pr olyl r esidues dur ing collagen synt hesis r equir es all of
t he following EXCEPT:
(a) Fe2
+
(b) a specific amino acid sequence.
(c) ascor bic acid (d) succinat e.
24. Dipht her ia t oxin:
(a) r eleases incomplet e polypept ide chains fr om t he r ibosome.
(b) act ivat es t r anslocase.
(c) pr event s r elease fact or fr om r ecognizing t er minat ion signals.
(d) at t acks t he RNA of t he lar ge subunit .
25. Tar get ing a pr ot ein t o be degr aded wit hin pr ot easomes usually r equir es ubiquit in. In
t he funct ions of ubiquit in all of t he following ar e t r ue EXCEPT:
(a) ATP is r equir ed for act ivat ion of ubiquit in.
(b) linkage of a pr ot ein t o ubiquit in does not always mar k it for degr adat ion.
(c) t he ident it y of t he N-t er minal amino acid is one det er minant of select ion for
degr adat ion.
(d) ATP is r equir ed by t he enzyme t hat t r ansfer s t he ubiquit in t o t he pr ot ein t o be
degr aded.
26. In all enzymes t he act ive sit e:
(a) cont ains t he subst r at e-binding sit e.
(b) is cont iguous wit h t he subst r at e-binding sit e in t he pr imar y sequence.
(c) cont ains a met al ion as a pr ost het ic gr oup.
(d) cont ains t he amino acid side chains involved in cat alyzing t he r eact ion.
27. Enzymes may be specific wit h r espect t o all of t he EXCEPT:
(a) chemical ident it y of t he subst r at e.
(b) t he at omic mass of t he element s in t he r eact ive gr oup (e.g., 12 C but not 14 C).
(c) Opt ical act ivit y of pr oduct for med fr om a symmet r ical subst r at e.
(d) which of a pair of opt ical isomer s will r eact .
28. The t r anspor t syst em t hat maint ains t he Na
+
a nd K
+
gr adient s acr oss t he plasma
membr ane of cells:
(a) involves an enzyme t hat is an ATPase.
(b) moves Na
+
eit her int o or out of t he cell.
(c) is an elect r ically neut r al syst em.
(d) in t he membr ane, hydr olyzes ATP independent ly of t he movement of Na
+
and
K
+
.
29. A mediat ed t r anspor t syst em would be expect ed t o:
(a) show a cont inuously incr easing init ial r at e of t r anspor t wit h in incr easing subst r at e
concent r at ion.
(b) exhibit st r uct ur al and/or st er eospecificit y for t he subst ance t r anspor t ed.
Practice Test Paper–Biochemistry 131
(c) be slower t han t hat of a simple diffusion syst em.
(d) est ablish a concent r at ion gr adient acr oss t he membr ane.
30. The gr oup t r anslocat ion t ype of t r anspor t syst em:
(a) does not r equir e met abolic ener gy.
(b) involves t he t r anspor t of t wo differ ent solut e molecules simult aneously.
(c) has been demonst r at ed for fat t y acids.
(d) r esult s in t he alt er at ion of t he subst r at e molecule dur ing t he t r anspor t pr ocess.
31. AIP-binding casset t e (ABC) t r anspor t er s:
(a) all have bot h a membr ane-spanning domain t hat r ecombine subst r at e and an
ATP-binding domain.
(b) all effect t r anslocat ion by for ming channels.
(c) all have t wo funct ions-for ming a channel and con r egulat ion.
(d) ar e all P-glycopr ot eins.
32. The mit ochondr ial membr ane cont ains a t r anspor t er for :
(a) NADH. (b) acet yl CoA.
(c) GTP. (d) ATP.
33. If r ot enone is added t o t he mit ochondr ial elect r on t r anspor t chain:
(a) t he P/O r at io of NADH is r educed fr om 3: 1 t o 2: 1
(b) t he r at e of NADH oxidat ion is diminished t o t wo-t hir ds of it s init ial value.
(c) succinat e oxidat ion r emains nor mal.
(d) elect r on flow is inhibit ed at sit e II.
34. If cyanide is added t o t ight ly coupled mit ochondr ia t hat ar e act ively oxidizing succinat e:
(a) subsequent addit ion of 2,4-dinit r ophenol will cause ATP hydr olysis.
(b) subsequent addit ion of 2,4-dinit r ophenol will r est or e succinat e oxidat ion.
(c) elect r on flow will cease, but ATP synt hesis will cont inue.
(d) subsequent addit ion of 2,4-dinit r ophenol and t he phosphor ylat ion inhibit or ,
oligomycin, will cause ATP hydr olysis.
35. The chemiosmot ic hypot hesis involves all of t he following EXCEPT:
(a) a membr ane imper meable t o pr ot ons.
(b) el ect r on t r a n s por t by t h e r es pi r a t or y ch a i n pu mps pr ot on s ou t of t h e
mit ochondr ion.
(c) pr ot on flow int o t he mit ochondr ia depends on t he pr esence of ADP and Pi’
(d) only pr ot on t r anspor t is st r ict ly r egulat ed; ot her posit ively char ged ions can
diffuse fr eely acr oss t he mit ochondr ial membr ane.
36. Suppose t he specific defect wer e a mut ant pyr uvat e dehydr ogenase (t he fir st cat alyt ic
subunit ) wit h poor binding of it s pr ost het ic gr oup. In t his t ype of defect , somet imes
gr eat ly incr easing t he diet ar y pr ecur sor of t he pr ost het ic gr oup is helpful. In t his
case incr easing which of t he following might be helpful?
(a) t hiamine(TPP) (b) niacin (for NAD)
(c) pant ot henic acid (for CoA) (d) r iboflavin (for FAD)
132 CSIR-NET Life Sciences
37. In glycolysis ATP synt hesis is cat alyzed by:
(a) hexokinase.
(b) 6-phosphofr uct o-I-kinase.
(c) glycer aldehyde 3-phosphat e dehydr ogenase.
(d) phosphoglycer at e kinase.
38. The ir r ever sible r eact ions of glycolysis include t hat cat alyzed by:
(a) phosphoglucose isomer ase.
(b) 6-phosphofr uct o-l-kinase.
(c) fr uct ose bisphosphat e aldolase.
(d) glycer aldehyde 3-phosphat e dehydr ogenase.
39. Glucokinase:
(a) has a Km consider ably gr eat er t han t he nor mal blood glucose concent r at ion.
(b) is inhibit ed by glucose 6-phosphat e.
(c) is also known as t he GLUT-2 pr ot ein.
(d) has glucose 6-phosphat ase act ivit y as well as kinase act ivit y.
40. In t he Cor i cycle:
(a) only t issues wit h aer obic met abolism (i.e., mit ochondr ial O
2
) ar e involved.
(b) a t hr ee-car bon compound ar ising fr om glycolysis is conver t ed t o glucose at t he
expense of ener gy fr om fat t y acid oxidat ion.
(c) glucose is conver t ed t o pyr uvat e in anaer obic t issues t his pyr uvat e r et ur ns t o
t he liver , wher e it is conver t ed int o glucose.
(d) t he same amount of ATP is used in t he liver t o synt hesis glucose as it is r eleased
dur ing glycolysis, leading t o no net loss on whole-body ener gy balance.
41. Gluconeogenic enzymes include all of t he following EXCEPT
(a) glucose 6-phosphat ase.
(b) phosphoenolpyr uvat e car boxykinase.
(c) phosphoglucomut ase.
(d) pyr uvat e car boxylase.
42. Glycosaminoglycans:
(a) ar e t he car bohydr at e por t ion of glycopr ot eins,
(b) cont ain lar ge segment s of a r epeat ing unit t ypically consist ing of a hexosamine
and a ur onic acid.
(c) always cont ain sulfat e.
(d) exist in only t wo for ms.
43. All of t he following st at ement s about acet yl-CoA car boxylase ar e cor r ect EXCEPT:
(a) i t u n der goes pr ot omer -pol ymer i n t er con ver s i on du r i n g i t s ph ys i ol ogi ca l
r egulat ion.
(b) it is inhibit ed by cAMP-mediat ed phosphor ylat ion.
(c) it is act ivat ed by bot h palmit oyl CoA and cit r at e.
(d) it s cont ent in a cell r esponds t o changes in fat cont ent in t he diet .
Practice Test Paper–Biochemistry 133
44. Pr imar y bile acids:
(a) ar e any bile acids t hat ar e found in t he int est inal t r act .
(b) ar e any bile acids r eabsor bed fr om t he int est inal t r act .
(c) ar e synt hesized in hepat ocyt es dir ect ly fr om cholest er ol.
(d) ar e conver t ed t o secondar y bile acids by conjugat ion wit h glycine or t aur ine.
45. St r uct ur al feat ur es t hat ar e common t o all pr ost aglandins include:
(a) 20-car bon at oms.
(b) an oxygen-cont aining int er nal het er ocyclic r ing.
(c) a per oxide gr oup at C-15.
(d) t wo double bonds.
46. Amino acids consider ed nonessent ial for humans ar e:
(a) t hose not incor por at ed int o pr ot ein.
(b) not necessar y in t he diet if sufficient amount s of pr ecur sor s ar e pr esent .
(c) t he same for adult s as for childr en.
(d) gener ally not pr ovided by t he or dinar y diet .
47. Aminot r ansfer ases:
(a) usually r equir e a-ket oglut ar amat e or glut amine as one of t he r eact ing pair .
(b) cat alyze r eact ions t hat r esult in a net use or pr oduct ion of amino acids.
(c) cat alyze ir r ever sible r eact ions.
(d) r equir e pyr idoxal phosphat e as an essent ial cofact or for t he r eact ion.
48. The amide nit r ogen of glut amine:
(a) r epr esent s a nont oxic t r anspor t for m of ammonia.
(b) is a major sour ce of ammonia for ur inar y excr et ion.
(c) is used in t he synt hesis of aspar agine. pur ines. and pyr imidines.
(d) can be r ecover ed as ammonia by t he act ion of glut aminase.
49. S-Adenosylmet hionine:
(a) cont ains a posit ively char ged sulfur (sulfonium) t hat facilit at es t he t r ansfer of
subst it uent s t o suit able accept or s.
(b) yields a-ket obut yr at e in t he r eact ion in which t he met hyl is t r ansfer r ed.
(c) donat es a met hyl gr oup in a fr eely r ever sible r eact ion.
(d) gener at es H
2
S by t r anssulfur at ion.
50. In humans, sulfur of cyst eine may par t icipat e in all of t he following EXCEPT:
(a) t he conver sion of cyanide t o less t oxic t hiocyanat e.
(b) t he for mat ion of t hiosulfat e.
(c) t he for mat ion of t aur ine.
(d) t he donat ion of t he sulfur for met hionine for mat ion.
134 CSIR-NET Life Sciences
51. Mor e fr ee ener gy is r eleased dur ing t he cit r ic acid cycle t han dur ing glycolysis, but
only 1 mole of ATP is pr oduced for each mole of acet yl CoA t hat ent er s t he cycle.
What happens t o most of t ile r emaining fr ee ener gy t hat is pr oduced dur il1g t he
cit r ic acid cycle?
(a) It is used t o synt hesize GTP
(b) It is used t o r educe elect r on car r ier s
(c) It is lost as heat
(d) It is used t o r educe pyr uvat e
52. The r educt ion of pyr uvat e t o lact ic acid dur ing fer ment at ion allows glycolysis t o
cont inue in t he absence of oxygen. Why?
(a) Wat er is for med dur ing t his r eact ion
(b) This r eact ion is a kinase r eact ion
(c) This r eact ion is coupled t o t he oxidat ion of NADH t o NAD
+
(d) This r eact ion is coupled t o t he r educt ion of NAD
+
t o NADH
53. Assume t hat eukar yot ic cell has abundant glucose and O
2
but needs ATP. The pr ot on
gr adient in mit ochondr ia of t he cell will be gener at ed by_____________ and used
pr imar ily for ––––––––––––––.
(a) t he elect r on t r anspor t chain; ATP synt hesis
(b) t he elect r on t r anspor t chain; subst r at e-level phosphor ylat ion
(c) glycolysis; pr oduct ion of H
2
O
(d) fer ment at ion; NAD r educt ion
54. The gener al name for an enzyme t hat t r ansfer s phosphat e gr oups fr om ATP t o a
Pr ot ein is
(a) Pr ot ein kinase (b) Phosphor ylase
(c) Phosphat ase (d) ATPase
55. All of t he following ar c pot ent ial cont r ol mechanisms for r egulat ion of gene expr ession
in eukar yot ic or ganisms EXCEPT
(a) Gene amplificat ion (b) The degr adat ion of mRNA
(c) The lact ose oper on (d) Tr anscr ipt ion
56. It is t heor et ically possible for a gene fr om any or ganism t o funct ion in any ot her
or ganism. Why is t his possible?
(a) All or ganisms have similar nuclei
(b) All or ganisms have t he same genet ic code
(c) All or ganisms ar e made up of cells
(d) All or ganisms have t r ansfer RNA
57. DNA fr agment s fr om a gel ar e t r ansfer r ed t o a membr ane via a pr ocedur e called
Sout her n blot t ing. The pur pose of Sout her n blot t ing is t o
(a) Analyze t he RFLPs in t he DNA
(b) Separ at e out t he PCRs’
(c) Per manent ly at t ach t he DNA fr agment s t o a subst r at e
(d) Separ at e t he t wo complement ar y DNA st r ands
Practice Test Paper–Biochemistry 135
58. 5'-phosphor ibosyl-1-pyr ophosphat e’(PAPP) is an int er mediat e in
(a) t he de nevo synt hesis of pur ine nucleot ides
(b) t he de novo synt hesis of pyr imidine nueleot ides
(c) t he salvage pat hway for t he synt hesis of pur ine nucleot ides
(d) All of t he above
59. In Lesch-Nyhan Syndr ome lack of act ivit y of t he defect ive enzyme (HGPRTase) should
r esult in higher t han nor mal t issue concent r at ions of all of t he following EXCEPT
(a) adenine (b) guanine
(c) ur ic acid (d) hypoxant hine
60. A comp l e t e l a ck of a d e n os i n e d e a mi n a s e ca u s e s SCI D (Se ve r e combi n e d
immunodeficiency). Which of t he following is LEAST t r ue
(a) Loss of t he enzyme causes incr eased levels of dATP because t her e is less t ur nover
of adenosine nucleosides in gener al
(b) Incr eased dATP decr eases t he concent r at ion of all r NTPs, blocking RNA synt hesis
(c) Incr eased dATP inhibit s r ibonucleot ide r educt ase, such t hat de novo pr oduct ion
of all r NDPs is inhibit ed
(d) Adenosine deaminase loss causes SCID because T cells ar e par t icular ly sensit ive
t o DNA r eplicat ion inhibit ion
61. DNA and RNA synt hesis polymer izat ion [of deoxynucleot ides] which t akes place
(a) In a 3' t o 5' dir ect ion
(b) In a 5' t o 3' dir ect ion
(c) In eit her (or bot h)dir ect ions
(d) DNA in 5' t o 3' and RNA in 3' t o 5'
62. TATA boxes and Pr ibnow boxes ar e component s of
(a) Oper at or s (b) Pr omot er s
(c) Enhancer s (d) Act ivat or s
63. The RNA in t he cell wit h t he gr eat est sequence diver sit y is
(a) Messenger RNA (b) Ribosomal RNA
(c) Tr ansfer RNA (d) (a) and (c)
64. Dur ing t he over all pr ocess of pr ot ein synt hesis, amino acids become covalent ly at t ached
t o
(a) Messenger RNA (b) Ribosomal RNA
(c) Tr ansfer RNA (d) Mor e t han one of t he above
65. Pr ot eins whose binding t o DNA act s t o pr event t r anscr ipt ion ar e known as
(a) Act ivat or s (b) Oper at or s
(c) Repr essor s (d) Tr anscr ipt ion fact or s
66. A common t ar get for ant ibiot ics in bact er ia is
(a) Micr osomes (b) Mesosomes
(c) Ribosomes (d) None of t he above
136 CSIR-NET Life Sciences
67. A piece of DNA of lengt h 10 KB obt ained using EcoRI digest ion is being inser t ed in a
pBR322 plasmid (appr oximat ely 4 KB) vect or which was fir st digest ed by t he r est r ict ion
enzyme EcoRI. The opt imum vect or : inser t r at io (µg/µg) should be appr oximat ely
(a) 0.1 (b) 0.5
(c) 1 (d) 5
68. Dr ied fish t r eat ed wit h salt and nit r at es has been found t o cont ain t he mut agen
2-chlor o-4-met hylt hiobut anoic acid (CMBA). Fr om what amino acid is CMBA der ived?
(a) Lysine (b) Cyst eine
(c) Met hionine (d) Valine
69. In exper iment s designed t o explor e whet her t her e is pr ot ein t r ansver se asymmet r y
in membr anes, it was shown t hat when t r ypsin was used t o t r eat int act er yt hr ocyt es,
t he car bohydr at e gr oups of t he t r ansmembr ane pr ot ein glycophor in was r eleased
fr om t he N-t er minus of t he pr ot ein (as glycopept ides). This was t aken as evidence
t hat t he N-t er minus of t he glycophor in must be on t he ext er ior side of t he membr ane
because
(a) t r ypsin is t oo lar ge t o get inside t he cell t o at t ack int er ior par t s of glycophor in.
(b) t r ypsin only at t acks car bohydr at e r esidues, not pr ot eins
(c) t r ypsin always at t acks near t he N-t er minus of a pr ot ein and never near t he
C-t er minus
(d) t r ypsin specifically cleaves pr ot eins at lysine or ar ginine r esidues
70. When a muscle cont r act s, what is happening t o t he Ca
++
levels inside and out side t he
cell?
(a) High amount s of cyt osolic Ca
++
ar e r eleased t o t he ext r acellular space
(b) Ion channels open t o allow ext r acellular Ca
++
t o flow int o t he cell
(c) Ca
++
fr om t he nucleus is r eleased t o t he cyt oplasm and t his t r igger s cont r act ion.
(d) Ca
++
ions at t ach st oma and t his causes muscle cont r act ion
71. Acet yl CoA ent er s t he TCA cycle by combining wit h:
(a) oxaloacet at e. (b) succinat e.
(c) cit r at e. (d) alpha-ket oglut er at e.
72. Cyt ochr ome c is a moveable per ipher al pr ot eins t hat is oxidized by complex:
(a) I . (b) II .
(c) III . (d) IV.
73. Which of t hese complexes wor k by r ot at ional cat alysis?
(a) NADH dehydr ogenase (complex I)
(b) Cyt ochr ome c oxidase (complex IV)
(c) Glycer ol phosphat e dehydr ogenase (glycer ol phosphat e shut t le)
(d) ATP synt hase
74. In which of t he following ways ar e per oxisomes NOT similar t o mit ochondr ia?
(a) Engages in oxidat ive met abolism.
(b) Has DNA t hat encodes a few of it s genes.
Practice Test Paper–Biochemistry 137
(c) Gener at ed by split t ing fr om pr e-exist ing or ganelles.
(d) Impor t s pr efor med pr ot eins fr om t he cyt osol.
75. Complex III (cyt ochr ome bc
1
) r eceives elect r ons dir ect ly fr om __________, which is a
lipid soluble membr ane molecule.
(a) ubiquinone (b) cyt ochr ome c
(c) NADH
+
(d) FADH
2
76. Wher e ar e ATP synt hase complexes found in t he mit ochondr ion?
(a) On t he out er membr ane. (b) In t he int er membr ane space.
(c) At t ached t o t he cr ist ae. (d) Float ing in t he mat r ix.
77. ATP synt hase is powed by a (n):
(a) pr ot on mot ive for ce. (b) elect r on gr adient .
(c) K
+
gr adient . (d) ATP.
78. Bot h t he mit ochondr ion and t he chlor oplast have all of t he following EXCEPT
(a) an inner and an out er membr ane. (b) ATP synt hase.
(c) cr ist ae. (d) elect r on t r anspor t .
79. Which of t hese molecules or complexes has a gr eat er oxidizing pot ent ial t han oxygen?
(a) Phot osyst em I (P700
+
) (b) Phot osyst em II (P680
+
)
(c) NADP
+
(d) plast iquinone
80. In t he Calvin cycle, CO
2
is fixed t o r ibulose bisphosphat e cat alyzed by an enzyme
familiar ly known as
(a) Ribozyme (b) LHCII
(c) Rubisco (d) P680
81. GLUT4 is an example of
(a) simple diffusion. (b) facilit at ed t r anspor t er .
(c) act ive t r anspor t er . (d) cot r anspor t er .
82. A hydr opat hy plot ident ifies pot ent ial membr ane spanning r egions of a pr ot ein by
analyzing t he
(a) pr imar y
(b) secondar y
(c) t er t iar y
(d) quat er nar y st r uct ur e of t he pr ot ein.
83. ABO blood t ypes ar e det er mined by a sacchar ide at t ached t o what ?
(a) Phosphoglycer ide (b) Sphingolipid
(c) Cholest er ol (d) Ion channel
84. Cholest er ol does what t o membr anes?
(a) Lower s t he t r ansit ion t emper at ur e.
(b) Accumulat es on t he out er leaf.
(c) Makes t he membr ane mor e hydr ophilic.
(d) Blur s t he t r ansit ion bet ween gel st at e and fluid st at e.
138 CSIR-NET Life Sciences
85. Which of t hese int egr al pr ot eins is an act ive t r anspor t er ?
(a) Kv cha nnel (b) GLUT4
(c) Na/K ATPase (d) aquapor in
86. Acet ocholine and nor epinephr ine ar e examples of:
(a) glycopr ot eins. (b) int egr al pr ot eins.
(c) ion channels. (d) neur ot r ansmit t er .
87. Epit helial cells wit hin t he int est ine can absor b glucose against a concent r at ion gr adient
by linking glucose t r anspor t wit h sodium t r anspor t , since sodium is much mor e
concent r at ed out side t he cell. What t er m best descr ibes t his st r at egy?
(a) Ant ipor t (b) Act ive t r anspor t
(c) Cot r anspor t (d) Impor t at ion
88. This most abundant ECM pr ot ein is for med of alpha-helix t r imer s t hat ar e bundled t o
for m r igid cable-like fibr ils.
(a) Collagen (b) Pr ot eoglycans
(c) Fibr onect in (d) Laminins
89. Which of t hese molecules definit ely indicat es t he pr esence of secondar y cell walls?
(a) Cellulose (b) Hemicellulose
(c) Pect in (d) Lignin
90. The concept t hat pr ot eins possess amino acids t hat act as an addr ess code is called
t he
(a) signal hypot hesis. (b) cell t heor y.
(c) pept ide t r ansfer ase pr oposal. (d) pr ot ein addr ess concept .
91. Which of t he following does NOT happen in cyclic phot ophosphor ylat ion?
(a) ATP is pr oduced
(b) Phot osyst em I r eact ion cent er is act ive
(c) Elect r on t r anspor t occur s in t he phot osynt het ic membr anes
(d) NADPH is for med
92. The conver sion of light ener gy t o chemical ener gy dur ing phot osynt hesis begins when
an excit ed pigment molecule:
(a) under goes fluor escence (b) loses ener gy as heat
(c) under goes an oxidat ion r eact ion (d) incr eases it s molecular mot ion
93. An over all r esult of phot osynt hesis in plant s is t he use of elect r ons fr om wat er t o
r educe:
(a) glucose (b) car bon dioxide
(c) oxygen (d) NADPH
94. Which of t he following is NOT t r ue of phot osyst em II?
(a) It is locat ed in t hylakoid membr anes.
(b) It is involved in t he oxidat ion of wat er .
(c) It has a special oxidizable chlor ophyll, P680.
(d) It is r equir ed for cyclic phot ophosphor ylat ion.
Practice Test Paper–Biochemistry 139
95. The elect r on t r anspor t chain is locat ed pr edominant ly in t he:
(a) Out er membr ane of t he mit ochondr ia
(b) Int er membr ane space of t he mit ochondr ia
(c) Inner membr ane of t he mit ochondr ia
(d) Mat r ix of t he mit ochondr ia
96. What cellular compar t ment becomes acidic (high concent r at ion of hydr ogen ions)
dur ing mit ochondr ial elect r on t r anspor t ?
(a) Mit ochondr ial st r oma
(b) Cyt oplasm
(c) Endoplasmic r et iculum
(d) Space bet ween inner and out er mit ochondr ial membr anes
97. In t he absence of oxygen, t he pr imar y pur pose of fer ment at ion is t o:
(a) pr oduce amino acids for pr ot ein synt hesis
(b) oxidize glucose t o gener at e r educe elect r on car r ier s
(c) gener at e alcohol for bever ages
(d) r egener at e NAD
+
fr om NADH allowing glycolysis t o cont inue
98. Which st at ement about enzyme cat alyzed r eact ions is NOT t r ue?
(a) enzymes for m complexes wit h t heir subst r at es.
(b) enzymes lower t he act ivat ion ener gy for chemical r eact ions.
(c) enzymes change t he K eq for chemical r eact ions.
(d) many enzymes change shape slight ly when subst r at e binds.
99. Act ivat ion ener gy is
(a) ener gy t hat must be added t o get a r eact ion st ar t ed, which is r ecover ed as t he
r eact ion pr oceeds
(b) differ ence in ener gy bet ween r eact ant s and pr oduct s
(c) ener gy t hat is lost as heat
(d) fr ee ener gy
100. Ener gy-r equir ing r eact ions can occur in biological syst ems because enzymes allow
t heir coupling t o ot her r eact ions wit h:
(a) an incr ease in ent r opy
(b) a low act ivat ion ener gy
(c) no inhibit or s
(d) pr oduct s of lower fr ee ener gy t han t he r eact ant s
3
Physiology
RESPONSE TO STRESS
St r e s s : St r ess can be defined as any envir onment al fact or t hat can have an adver se effect on a
Plant ’s gr owt h, r epr oduct ion, and sur vival. Plant s can r espond t o st r ess in sever al ways t hey
can escape t he effect s of st r ess by complet ing t heir gr owt h dur ing less st r essful per iods or t hey
ma y suffer injur y if t he st r ess is pr esent a nd t hey ca nnot cope. Pla nt s ca n cope wit h
envir onment al st r ess t hr ough a combinat ion of development and physiological r esponses.
Response to water stress
On br ight , war m, dr y day, a plant may be st r essed by a wat er deficiency because it is losing
wat er by t r anspir at ion fast er t han t he wat er can be r est or ed by upt ake fr om t he soil. This
cont inuous dr ought can st r ess plant s for weeks or mont h. Sever e wat er defict , may kill a plant .
But plant s have cont r ol syst ems t hat enable t hem t o cope wit h less ext r eme wat er defict s.
Many of a plant s r esponses t o wat er defict help t he plant conser ve wat er by r educing t he r at e
of t r anspir at ion. Wat er defict can st imulat es incr eased synt hesis and r elease of abscisic acid
fr om mesophyll cells in t he leaf, and t his hor mone helps keep st omat a closed by act ing on
guar d cell membr anes and causes guar d cell t o loose t ur gor . Leaves r espond t o wat er defict in
sever al ot her ways, because cell expansion is a t ur gor - dependent pr ocess, a wat er defict will
inhibit t he gr owt h (expansion) of yong leaves. This r esponse minimizes t he t r anscr ipt ional loss
of wat er by slowing t he incr ease in leaf sur face. When t he leaves of many gr asses and ot her
plant s wilt fr om a wat er defict , t hey r oll int o a shape t hat r educes t r anscr ipt ion by exposing
less leaf sur face t o t he sun. While all of t hese r esponses of leaves help t he plant conser ve
wat er , t hey also r educes phot osynt hesis. This is one r eason a dr ought diminished cr op yield.
Response To Oxygen Stress
An over wat er ed house plant may suffocat e because t he soil lacks t he air spaces t hat pr ovide
oxygen for cellular r espir at ion in t he r oot s. Some plant s ar e st r uct ur ally adapt ed t o ver y wet
habit at s. For example, t he submer ged r oot s of t r ees called mangr oves, which inhabit coast al
Physiology 141
mar shes, ar e cont inuous wit h aer ial r oot s t hat pr ovide access t o oxygen. Exper iment ally it was
shown t hat oxygen depr ivat ion st imulat es t he pr oduct ion of hor mone et hylene, which causes
some of t he cells in t he r oot s cor t ex t o age and die. Enzymat ic dest r uct ion of t he cell walls
cr eat es air t ubes t hat funct ion as ‘snor kes’ pr oviding oxygen t o t he submer ged r oot s.
Response To Salt Stress
An excess of sodium chlor ide or ot her salt s in t he soil t hr eat ens plant s for t wo r easons. Fir st ,
by lower ing t he wat er pot ent ial of t he soil solut ion, salt can cause a wat er defict in plant s even
t hough t he soil has plent y of wat er . This is because in an envir onment wit h a wat er pot ent ial
mor e negat ive t han t hat of t he r oot t issue, r oot s will lose wat er r at her t han absor b it . The
second pr oblem wit h saline soil is t hat sodium and cer t ain ot her ions ar e t oxic t o plant s when
t heir concent r at ion is r elat ively high. The select ively per meable membr anes of r oot cells impede
t he upt ake of most har mful ions, but t his only aggr avat es t he pr oblem of acquir ing wat er fr om
soil t hat is r ich in solut es. Many plant s can r espond t o moder at e soil salanit y by pr oducing
compat ible solut es, or ganic compounds t hat keeps t he wat er pot ent ial of cells mor e negat ive
t han t hat of t he soil solut ion wit hout admit t ing t oxic quant it ies of salt . However , most plant s
cannot sur vive salt st r ess for ver y long. The except ion ar e halophyt es, salt t oler ant plant s wit h
special adapt at ions such as salt glands, which pump salt s out of t he plant acr oss t he leaf
epider mis.
Response to Heat Stress
Exccesive heat can har m and event ually kill a plant by denat ur ing it s enzymes and damaging
it s met abolism in ot her ways. One funct ion of t r anspir at ion is evapor at ive cooling. On a war m
day, for example, t he t emper at ur e of a leaf is below air t emper at ur e which favour s t r anspir at ion
and causes wat er deficiency in many plant s; t he closing of st omat a in r esponse t o t his st r ess
conser ves wat er but sacr ifices evapor at ive cooling. This dilemma is one of t he r easons t hat
ver y hot , dr y days t ake such a t oll on most plant s. Many plant s r esponse t o t his st r ess by
synt hesizing r elat ively lar ge quant it ies of special pr ot eins called h e a t s h ock p r ot e i n s . Some
heat shock pr ot eins ar e ident ical t o chaper on pr ot eins, which funct ion in unst r essed cells as
t empor ar y scaffolds t hat ot her pr ot eins fold int o t heir funct ional confor mat ions.
Response to Cold Stress
One pr oblem plant s face when t he t emper at ur e of t he envir onment falls is a change in t he
fluidit y of cell membr anes. When a membr ane cools below a cr it ical point , it loses it s fluidit y as
t he lipids become locked int o cr yst alline st r uct ur es. This alt er s solut e t r anspor t acr oss t he
membr ane and also adver sely affect s t he funct ions of membr ane pr ot eins. Plant s r espond t o
cold st r ess by alt er ing t he lipid composit ion of t heir membr anes. For example, membr ane
lipids incr ease in t heir pr opor t ion of unsat ur at ed fat t y acids, which have shapes t hat help keep
membr anes fluid at lower t emper at ur e by impeding cr yst al for mat ion. Fr eezing is a mor e
sever e ver sion of cold st r ess. At fr eezing t emper at ur es, ice cr yst als begin t o for m in most
plant s. If ice cr yst als ar e confined t o cell walls and Int er cellular spaces, t he plant will pr obably
sur vive. However , if ice cr yst als begin t o for m wit hin pr ot oplast s, t he shar p cr yst al per for at e
membr anes and or ganelles, killing t he cells. However , plant s like oaks, maple, r hododendr ons
nat ive t o r egions wher e wint er s ar e cold have special adapt at ions t hat enable t o cope wit h
fr eezing st r ess. For example, changes in t he solut e composit ion of live cells allow t he cyt osol t o
cool below 0 degr ee Celsius wit hout cr yst al for mat ion Alt hough ice cr yst als may for m in t he
cell walls.
142 CSIR-NET Life Sciences
TRANSPORT ACROSS MEMBRANES
1. Move me n t of Sma l l Mol e cu l e s Acr os s Me mb r a n e s ca n i n vol ve s i mp l e
d i ffu s i on or p r ot e i n -me d i a t e d t r a n s p or t . Ce l l me mb r a n e s a r e s e l e ct i ve l y
p er mea ble.
(a) P a s s i ve Di ffu s i on . Lipophilic solut es cr oss t he membr ane fr eely by dissolving
in t he lipid bilayer . This is passive diffusion. Exa mp l e s : e t h a n ol (a lcohol,
cont ains bot h polar and non-polar r egions); also fa t t y a ci d s , glyce r ol, s t e r oi d s ,
et c. Also nonpolar gases like O=O (O
2
)
(b) Se l e ct i ve t r a n s p or t b y p r ot e i n ca r r i e r s = “p e r me a s e s ”. P ol a r or i on i c
small solut es may be t r anspor t ed acr oss membr anes if specific pr ot ein car r ier s
ar e in t he membr ane. Exa mp l e s : sugar s, amino acids, ions.
(c) Ma n y s u b s t a n ce s ca n n ot cr os s t h e me mb r a n e . Exa mp l e s : la r ge molecules
such as pr ot eins, nucleic acids. Also small polar molecules or ions for which
t her e is no pr ot ein car r ier .
2. Some p r ot ei n t r a n sp or t er s r eq u i r e en er gy; ot h er s d o n ot . Th er e a r e 2 p ossi b le
s i t u a t i on s :
(a) fa c i l i t a t e d d i ffu s i on . Membr ane has specific pr ot ein car r ier , will bind t o
molecule and br ing it acr oss cell membr ane. No ener gy r equir ed. No pr efer ent ial
dir ect ion. If molecule is mor e concent r at ed out side t han inside cell, net movement
will be out of cell. Examples include-
• Glucose t r anspor t er s- 5 differ ent GLUT pr ot eins and 2 t ypes t hat cot r anspor t
Na and glucose (t hese ar e used for secondar y act ive t r anspor t )
• Wat er channels- 8 differ ent t ypes of aquapor ins
(b) a ct i ve t r a n s p or t . Membr ane has specific pr ot ein car r ier , also a r equir ement
for ener gy (ATP or ot her for m of ener gy). Will move solut e against a concent r at ion
gr adient , so can concent r at e mat er ial even if diffusion would favor opposit e
dir ect ion of flow.
Exa mp l e: Na
+
- K
+
ATP a s e i n n er ve cel l s . Pu mps Na
+
t o ou t s i de, K
+
in,
ma int a ins elect r ica l pot ent ia l a ga inst diffusion. When ner ve cell “fir es”,
moment ar y gat es open t o let diffusion occur . Then pumps ar e t ur ned back on t o
r est or e pot ent ial.
3. Act i ve t r a n s p or t ca n i n vol ve ATP p u mp s , s ymp or t , or a n t i p or t
(a) ATP p u mp s .
• ATP-power ed pumps (ATPases) couple t he split t ing, or hydr olysis, of ATP
wit h t he movement of ions acr oss a membr ane against a concent r at ion
gr adient .
• ATP is hydr olyzed dir ect ly t o ADP and inor ganic phosphat e, and t he ener gy
r eleased is used t o move one or mor e ions acr oss t he cell membr ane.
• As much as 25% of a cell’s ATP r eser ves may be spent in such ion t r anspor t .
• Examples include:
1. The Na
+
-K
+
ATPase pumps Na
+
out of t he cell while it pumps K
+
in.
Because t he pump moves t hr ee Na
+
t o t he out side for ever y t wo K
+
t hat ar e moved t o t he inside, it cr eat es an over all char ge separ at ion
known as polar izat ion. This elect r ical pot ent ial is r equir ed for ner vous
Physiology 143
syst em act ivit y, and supplies ener gy needed for ot her t ypes of t r anspor t
such as sympor t and ant ipor t .
2. Ca
++
ATPases ar e r esponsible for keeping int r acellular Ca
++
at low
levels, a necessar y pr econdit ion for muscle cont r act ion.
(b) Symp or t
• To t r anspor t some subst ances against a concent r at ion gr adient , cells use
ener gy alr eady st or ed in ion gr adient s, such as pr ot on (H
+
) or sodium (Na
+
)
gr adient s, t o power membr ane pr ot eins called t r anspor t er s.
• When t he t r anspor t ed molecule and t he co-t r anspor t ed ion move in t he
same dir ect ion, t he pr ocess is known as sympor t .
• Example: t r anspor t of amino acids acr oss t he int est inal lining in t he human
gut .
(c) An t i p or t
Cell uses movement of an ion acr oss a membr ane and down it s concent r at ion
gr adient t o power t he t r anspor t of a second subst ance “uphill” against it s gr adient .
• In t his pr ocess, t he t wo subst ances move acr oss t he membr ane in opposit e
dir ect ions.
• Example: t r anspor t of Ca
2+
ions out of car diac muscle cells. Muscle cells
ar e t r igger ed t o cont r act by a r ise in int r acellular Ca
2+
concent r at ion, so it
is imper at ive t hat Ca
2+
be r emoved fr om t he cyt oplasm so t hat t he muscle
can r elax befor e cont r act ing again. This ant ipor t syst em is so effect ive t hat
it can maint ain t he cellular concent r at ion of Ca
2+
at levels 10,000 t imes
lower t han t he ext er nal concent r at ion.
4. Move me n t of La r ge Mol e cu l e s : Exocyt os i s & En d ocyt os i s
(a) Lar ge molecules (pr ot eins, nucleic acids, polypept ides lar ger t han a few amino
acids, polysacchar ides lar ger t han a few sugar s) ar e not car r ied by t r anspor t
pr ot eins.
(b) Ther e ar e mechanisms for moving lar ger molecules, but t hey don’t ent er int o
cyt oplasm.
(i) Exocyt os i s : membr ane vesicle fuses wit h cell membr ane, r eleases enclosed
mat er ial t o ext r acellular space. Ex: r elease of digest ive enzymes fr om
pancr eat ic cells; mucus, milk, hor mones, et c.
(ii) E n d o c y t o s i s: cell membr a ne inva gina t es, pinches in, cr ea t es vesicle
enclosing cont ent s. Thr ee common sit uat ions:
1. P h a gocyt os i s : Typically wor ks on debr is, bact er ia, ot her par t iculat e
mat t er . Cont ent s of t he “phagosome” ar e usually fused wit h lysosome
t o cr eat e “phagolysosome”, wher e mat er ial is br oken down. Especially
common i n wh i t e bl ood cel l s s u ch a s ma cr oph a ges a n d ot h er
leukocyt es.
2. P i n ocyt os i s : similar t o phagocyt osis, but ingest s fluid r at her t han
par t iculat e mat t er . “Cell dr inking”. Ex. cells lining blood capillar ies
t a ke fluid fr om blood (but not r ed cells), move fluid a cr oss t heir
cyt oplasm, r elease int o ext r acellular space sur r ounding cells out side
t he capillar y.
144 CSIR-NET Life Sciences
3. Ca r r i e r -m e d i a t e d e n d o c y t o s i s ( CME ) / r e c e p t o r -m e d i a t e d
e n d ocyt os i s : ver y specialized syst em. Cer t ain impor t ant molecules
or ions ar e not br ought int o cell by t r anspor t pr ocesses, but by CME.
The body uses hor mones t o r egulat e membr ane t r anspor t in t his way
Eg. 1. ir on is car r ied t hr ough blood t ight ly bound t o t r ansfer r in pr ot ein
car r ier . To get ir on int o cells, cell membr ane cont ains special r ecept or
pr ot eins t ha t bind t r a nsfer r in, move t owa r ds specia l r egions of
membr ane under which lie cla t h r i n pr ot eins. Endocyt osis occur s inside
clat hr in “cage”, moves inside cell. Cage event ually r ecycles back t o
cell sur face, r et ur ning t r ansfer r in pr ot eins t o cell ext er ior . However ,
ir on is r eleased inside cell, exit s fr om vesicles, becomes bound t o
fer r it in.
5. Ce l l s Re gu l a t e P e r me a b i l i t y b y Ad d i n g & Re movi n g Me mb r a n e Tr a n s p or t
P r ot e i n s
(a) Insulin is t r anspor t ed by t r anspor t er GLUT4. Insulin causes glucose t r anspor t
molecules t o be inser t ed int o muscle and adipose t issue cells. Glucose is t hen
t aken up int o t hose t issues, lower ing t he blood concent r at ion.
(b) Ant idiur et ic Hor mone (ADH) is t r anspor t ed via aquapor in. ADH causes aquapor in
2 pr ot eins t o be added t o t he kidney collect ing duct membr anes. The r esult is
wat er conser vat ion.
(c) Aldost er one is t r anspor t ed by Na Pump. Aldost er one causes cells in t he dist al
t ubules and collect ing duct of t he kidney t o make mor e Na pump molecules.
The final r esult is t hat t he body r et ains mor e Na and secr et es mor e K int o t he
ur ine.
PLANT HORMONES
In all plant s, t her e occur in minut e quant it ies of cer t ain subst ances (plant gr owt h r egulat or s or
phyt ohor mones) which r egula t e gr owt h a nd differ ent ia t ion. Five ma jor t ypes of gr owt h
subst ances ar e r ecognized: auxins and gibber ellins (bot h concer ned wit h cell enlar gement and
differ ent iat ion); cyt okinins (concer ned wit h cell division); abscisic acid (wit h r est ing st at es like
lat er al buds); and et hylene (wit h senescence-ageing),
1. Auxins
The auxins ar e weak or ganic acids wit h t he acidic gr oup posit ioned at t he end of t he side chain
at t ached t o an unsat ur at ed r ing(s) syst em. Pr esent ly, indole acet ic acid (IAA) and ot her nat ur al
or synt het ic gr owt h-r egulat ing subst ances having st r uct ur es and funct ions similar t o IAA, ar e
t er med auxins.
(i) Cha r les Da r win conduct ed his exper iment s concer ning gr owt h on ca na r y gr a ss
(Phalaris canariensis ).
(ii) Went is cr edit ed wit h t he discover y of auxin.
(iii) Auxins ar e synt hesized mainly in apices and exhibit polar t r anspor t .
(iv) Using Avena cur vat ur e t est , Kogl and Haagen Smit (1931) found t hat human ur ine
cont a ined a gr owt h subst a nce, which wa s isola t ed a nd given t he na me a uxin a
(auxent r iolic acid). In 1934, Kogl and cowor ker s isolat ed anot her compound auxin-b
Physiology 145
(auxenolinic acid) fr om cor n ger m oil and het er oauxin (now known as IAA or indole 3-
acet ic acid) fr om human ur ine. The auxins a and b could not be isolat ed again and
t heir exist ence is doubt ful but IAA has been isolat ed in cr yst alline for m fr om differ ent
sour ces by differ ent invest igat or s at differ ent t imes.
(v) Pla nt s ha ving excur r ent ha bit (pa lms, conifer s, polya t hia ) ha ve gr ea t er a pica l
dominance.
Functions of auxins
(i) Cell elongat ion
(ii) Cell divisions
(iii) Phot ot r opism
(iv) Geot r opism
(v) Apical dominance
(vi) Root init iat ion
(vii) Abscission
(viii) Feminising effect
(ix) Par t henocar py
(x) Weedicide
(xi) Pr event ion of lodging
Applications of synthetic auxins
1. Root i n g: IBA, IBA-alanine and NAA ar e used.
2. P a r t h e n oca r p y: IAA, IBA.
3. Weed i ci d e: 2,4-D, 2,4, 5- T.
4. F l owe r i n g: NAA and 2,4-D for lit chi and pineapple.
5. St or a ge : Met hyl est er of NAA for st or age of pot at o.
6. P r e h a r ve s t fr u i t d r op : 2,4-D for cit r us fr uit s, NAA for t omat o.
7. P r e ve n t i on of l od gi n g: Napht halene acet amide.
8. Veget a b le cr op s: Chlor ophenoxypr opionic acid is used t o impr ove qualit y of veget able
cr ops.
9. Dwa r f s h oot s : NAA for incr easing dwar f shoot s and number of fr uit s in apple.
2. Gibberellins
Gibber ellins ar e synt hesized in t he apices of young leaves and r oot s and ar e t r anspor t ed t hr ough
xylem. Ant i-giber ellins ar e synt het ic compounds which int er fer e wit h t he synt hesis of gibber ellins
in t he pla nt body, hence ca lled gr owt h r et a r da nt s e.g. ma leic hydr a zide, phosphon D,
chlor ocholine chlor ide (CCC).
J apanese far mer not iced ‘bakanae’ or ‘foolish seedling disease of r ice’. As a r esult of t he
disease, cer t ain r ice seed lings gr ew excessively t all and r apidly; and t oppled over befor e for ming
seeds. Cr op losses as high as 40% wer e r epor t ed. In t he beginning of t went iet h cent ur y, it
became known t hat t he disease is caused by a fungus Gibberella fujikuroi (per fect st at e of
Fuasarium moniliforme).
146 CSIR-NET Life Sciences
Kur osowa (1926) showed t hat t he st er ile filt r at es of t he fungus ar e capable of causing t he
sympt oms of bakanae disease in ot her wise nor mal seedlings. Yabut a and Hayashi (1910) isolat ed
t he gr owt h inducing pr inciple and called it Gibber ellin. In 1938, Yabut a and Sumiki wer e able
t o isolat e t he cr yst alline for m of gibber ellin fr om t he cult ur e of t he fungus. This and ot her
similar compounds isolat ed wer e called gibber ellins (aft er t he name of t he fungus Gibber ella)
and wer e number ed GA
1
, GA
2
, GA
3
.
Chemically all gibber ellins ar c t er penes a complex gr oup of plant chemicals r elat ed t o
lipids. All ar e weak acids. Gibber ellins ar e found in abundance in young expanding or gans
being synt hesized especially in embr yos, young apical leaves, buds, seeds and r oot t ips. Aft er
synt hesis, t hese ar e t r anslocat ed up or down t he plant s fr om t he leaves.
Applications of gibberellins
(i ) I n t e r n od a l e l on ga t i on : Like auxins, t he main effect of gibber ellins is on st em
elongat ion mainly by affect ing cell elongat ion. Gibber ellins st imulat e st em elongat ion
and leaf expansion, but do not affect r oot s. Thus, gibber ellins r est or e nor mal size and
gr owt h t o genet ically dwar f var iet ies of pear and maize. Though st em gr owt h of
nor mal plant s is also pr omot ed, but her e t hese do not induce mar ked elongat ion. It is
believed t hat cer t ain specific t ypes of dwar fness in plant s ar e due t o deficiency of
gibber ellin.
(i i ) Bol t i n g: In many plant s, leaf development is pr ofuse, while int er node gr owt h is
r et a r ded. This for m of gr owt h is ca lled “r oset t e” e.g., ca bba ge. J ust befor e t he
r epr oduct ive phase, t he int er nodes elongat e enor mously causing a mar ked incr ease
in height . The st em somet imes elongat es fr om 5-6 t imes t he or iginal height of t he
plant . This is called bolt ing. Bolt ing r equir es eit her long days or cold night s. So if a
cabbage head is kept under war m night s, it r et ur ns t o it s r oset t e habit . However ,
applicat ion of gibber ellins t o t hese plant s can induce bolt ing even under t he condit ions
t hat would nor mally favor r oset t e for m. The gibber ellins t hus appear t o have a r ole
in bolt ing or not bolt ing of a plant . Some gibber ellin-like subst ances found in gr eat er
amount s in bolt ed plant s t han in t he non-bolt ed for m.
(i i i ) Ge r mi n a t i on of s e e d s : especially in cer eals, is t r igger ed by soaking t he seed in
wat er . Aft er imbibit ion of wat er , t he embr yo secr et es gibber ellin which diffuses t o
t he aleur one layer , st imulat ing t he synt hesis of sever al enzymes, including amylase,
pr ot eases, lipases. These enzymes cat alyse t he br eakdown of food r eser ves in t he
endosper m and t he pr oduct s liber at ed diffuse t o t he embr yo, wher e t hey ar e used in
gr owt h.
(v) Con t r ol of fl owe r i n g: Gibber ellins pr omot e flower ing in long day plant s and inhibit
it in shor t day plant s. These also cont r ol sex-expr ession in cer t ain species. In gener al,
t he applicat ion of gibber ellins pr omot es t he pr oduct ion of male flower s in female
plant s of Callilabis.
(vi ) Con t r ol of fr u i t gr owt h : Along wit h gibber ellins, t he auxins cont r ol fr uit gr owt h
and development . Gibber ellins cause par t henocar py in pome fr uit s (apple, pear et c.)
and ar e now-a-days used ext ensively t o incr ease t he fr uit size and bunch lengt h in
gr ape.
(vi i ) Ve r n a l i za t i on : Gibber ellins can subst it ut e ver nalizat ion.
(vi i i ) Dor ma n cy: Gibber ellins over come nat ur al dor mancy of buds, t uber s, seeds et c.
Physiology 147
Commercial application
1. F r u i t gr owt h – incr ease number and size of gr apes, t omat o.
2. P a r t h e n oca r p y – in pomes.
3. Ma l t - Incr ease yield of mall fr om bar ley.
4. De l a ye d r i p e n i n g – in cit r us.
5. Ove r comi n g d or ma n cy – in phot oblaslic seeds of t obacco and let t uce.
6. F l owe r i n g – in long day plant s in non-induct ive per iod.
3. Cytokinins
Cyt okinins ar e subst ances which act pr imar ily on cell division and have lit t le or no effect on
ext ension gr owt h. A number of compounds capable of pr omot ing cell division ar e found in
plant s.
Dur ing t he per iod 1954-1956, Skoog found t hat coconut milk cont ained it subst ance t hat
st imulat ed cell division in t obacco pit h cult ur es. In 1955, Miller et . al. separ at ed fr om yeast
DNA an act ive st imulant of cell division which t hey called kinet in (because of it s involvement
in cell division i.e. cyt okinesis). Lat er on, t he subst ance was ident ified as 6-fur fur yl aminopur ine.
Subsequent ly, t he t er m cyt okinin was adopt ed.The fir st nat ur ally occur r ing cyt okinin t o be
chemically ident ified was fr om young maize (Zea mays) gr ains in 1963 and was called zeat in.
Applications of cytokinins
(i) Ce l l -d i vi s i on : Cyt okinins ar e quit e abundant wher ever , r apid cell division occur s,
especially in gr owing t issues such as embr yos, developing fr uit s and r oot s. In mat ur e
plant s, t hese ar e fr equent ly synt hesized in t he r oot s and move t o t he shoot s t hr ough
xylem.
(ii) Mor p h oge n e s i s : Cyt okinins pr omot e cell division. However , t hese never act alone.
In t he pr esence of auxins, cyt okinins pr omot e cell division even in non-mer ist emat ic
t issues. In t issue cult ur es of par enchyma, mit ot ic divisions ar e acceler at ed when
bot h auxin and cyt okinin ar e pr esent . However , no r esponse occur s wit h auxin or
cyt okinin alone. Also, t he r at io of cyt okinins t o auxins also cont r ols cell differ ent iat ion.
When bot h of t hese ar e pr esent in r elat ively equal quant it ies t he cells do divide but -
fail t o differ ent iat e. If t her e is mor e cyt okinin t han auxin, shoot buds develop.
Relat ively mor e auxin t han cyt okinins leads t o t he development s of r oot s.
(iii) Ap i ca l Domi n a n ce : Cyt okinin and auxin can act ant agonist ically in t he cont r ol of
apical dominance. While auxins pr omot e t he gr owt h of apical buds, t he cyt okinin
st imulat e t he gr owt h of lat er al buds.
(iv) Dela y i n sen escen ce: Cyt okinins delay t he senescence of plant or gans by cont r olling
pr ot ein synt hesis and mobilizat ion of r esour ces. This is evidenced by t he fact t he cut
leaves dipped in cyt okinins st ay gr een longer t han t he cont r ol leaves. This is called
Richmond Lang effect .
(v) F l owe r i n g: Cyt okinins also induce flower ing in cer t ain species of plant s and ar e also
r esponsible for br eaking t he dor mancy of seeds of some plant s
(vi) P h l oe m t r a n s p or t a n d Accu mu l a t i on of s a l t s i n t h e ce l l s .
(vii) Se x-e xp r e s s i on – pr omot e femaleness
148 CSIR-NET Life Sciences
Commercial application
1. Tissue cult ur e.
2. Shelf life of veget ables, flower s is incr eased.
3. Over coming senescence.
4. Ethylene
Et hylene is t he only gaseous nat ur al plant gr owt h r egulat or . Some of t he inhibit or y effect s
which wer e ear lier t hought t o be due t o auxins ar e now known t o be t he effect of et hylene.
A r ipe or injur ed fr uit in a cont ainer hast ens t he r ipening of ot her fr uit s. ‘This is ver y
common obser vat ion and belief. Mer chant s have been using ker osene lamps and hay for hast ening
t he development of color in fr uit s. These effect s ar e due t o et hylene. In 1930’s it became known
t hat et hylene gas hast ened t he r ipening of cit r us fr uit s and affect ed plant s gr owt h in differ ent
ways. Lat er on, it was found t hat cer t ain r ipe fr uit s e.g., bananas pr oduced a gas wit h similar
effect s. In 1934, mat ur ing apple was shown t o emit et hylene. Lat er on, it was found t hat
et hylene is emit t ed by a wide var iet y of r ipening fr uit s, and fr om wounded or gans.
Et hylene is pr oduced by most or all plant or gans. High concent r at ions of auxin induce t he
for mat ion of et hylene. Though it is a gas, it does not genenr ally move t hr ough t he air spaces in
t he plant s. Rat her , it escapes fr om t he plant s sur face.
Applications of ethylene
(i) Gr owt h : It inhibit s st em elongat ion and st imulat es it s t r ansver se expansion. As a
r esult , t he st em looks swollen.
(ii) Ab s ci s s i on : It acceler at es abscission of leaves, flower s and fr uit s.
(ii) F r u i t r i p e n i n g: It s chief effect s ar e on t he r ipening of fr uit s accompanied by a r ise
in t he r at e of r espir at ion (climact er ic).
(iv) F l owe r i n g: Applicat ion of et hylene induces flower ing in pineapp.12 and st imulat es
t he r ipening of cit r us fr uit s and t omat oes. It s applicat ion incr eases t he number of
female flower s and fr uit s in cucumber plant s.
A commer cial compounds ‘e t h e p h on ’ br eaks down t o r elease et hylene in plant s. It is
par t icular ly applied t o r ubber plant s t o st imulat e t he flow of lat ex.
5. Abscisic Acid
For a long t ime it has been suspect ed t hat dor mancy is caused by inhibit or s. A gr oup of scient ist s
led by War eing init iat ed st udies t o find t hem. In 1964, pur e cr yst als of a subst ance wer e called
dor min. It was found t o be similar t o anot her compound isolat ed fr om young cot t on fr uit s in
1963 by anot her gr oup of scient ist s. This subst ance acceler at ed abscission and was called
abscission II. In 1967 it was decided t o call it abscisic acid.(ABA)
Since t hen, it has been found in all gr oups of Plant s (fr om mosses t o higher plant s). In
liver wor t s and algae a compound lunular ic acid had been found having act ivit ies similar t o
ABA. Chemically, ABA is a t er per ioid. It is t he only gr owt h subst ance in it s class.
ABA is synt hesized in t he leaves, st ems, fr uit s and seeds. Isolat ed chlor oplast can synt hesize
it . It is dist r ibut ed t hr ough t he vascular syst em, especially phloem. ABA is a major inhibit or of
gr owt h in plant s and is ant agonist ic t o all t he t hr ee gr owt h pr omot er s.
Physiology 149
APPLICATIONS OF ABA
(i) St op p a ge of ca mb i a l a ct i vi t y – It inhibit s mit osis in vascular cambium.
(ii) Bu d d o r ma n c y – I t induces a xilla r y buds t o become dor ma nt a s t he wint er
appr oaches.
(iii) Se e d d or ma n cy – It induces dor mancy in seeds.
(iv) Tr a n s p i r a t i on – It is a ‘st r ess hor mone’ and helps t he plant t o cope wit h adver se
envir onment al condit ions especially dr ought . In wilt ing t omat o leaves, ABA br ings
about t he closur e of st omat a.
Commercial uses
1. It may be spr ayed on t r ee cr ops t o r egulat e fr uit dr op at t he end of t he season.
2. Applicat ion of ABA t o gr een or anges t ur ns t hem yellow by inducing synt hesis of
car et enoids.
3. Ant i-t r anspir ant .
ANIMAL HORMONES
1. Exocr i n e gl a n d s ar e t hose which dr ain out t heir secr et ion t hr ough a d u ct , e .g.
l i ve r , i n t e s t i n a l gl a n d s , ga s t r i c gl a n d s .
2. En d ocr i n e gl a n d s . The glands, which lack duct and dischar ge, t heir secr et ions int o
t he blood st r eam ar e called as Endocr ine glands. They ar e also called as d u ct l e s s
gl a n d s . The chemical secr et ions of endocr ine glands ar e called h or mon e s and ar e
popular ly called t he ch e mi ca l me s s e n ge r s of t he body. They affect definit e par t of
t he body and pr epar e it for a specific funct ion. The par t of t he body wher e a hor mone
pr oduces it s effect is called t he t ar get .
• En d ocr i n e s e cr e t i on . Tar get or gans lies at a dist ance or is far r emoved fr om
t he sit e of or igin of hor mone.
• P a r a cr i n e se cr e t i on . Act ing on cells in near vicinit y.
• Au t ocr i n e s e cr e t i on . Act ing on same cells, which secr et e it .
3. Me ch a n i s m of a ct i on of h or mon e s :
Pr ot ein hor mones act on t he t ar get r ecept or s pr esent on plasma membr ane. These
r ecept or s in t ur n excit e second messenger s inside t he nucleus for act ion on DNA for
differ ent ial t r anscr ipt ion second messenger can be.
• cAMP (3' 5' cyclic a denosine mono phospha t e) e.g. TSH, LH, ACTH, FSH,
vasopr essin, par at hyr oid hor mone, glucagons, secr et in, cat echolamines.
• Ca
++
ions and calmodulin (calcium binding pr ot ein) e.g. ?
• Inosit ol phosphat e and diacylglycer ol
4. The na me h or mon e was fir st used by t he English physiologist s. M. Ba yl i s s a n d
E.H. St a r l i n g in 1909. A hor mone (Gr . Hor mao = t o excit e) may be defined as a
specific or ganic pr oduct of an endocr ine gland secr et ed int o t he blood which car r ies it
t o some par t of t he body (t ar get or gan) wher e it r egulat es a definit e physiological
effect .
150 CSIR-NET Life Sciences
5. He t e r ocr i n e gla n d s ar e t hose glands which ar e exocr ine as well as endocr ine in
funct ions e.g. t est es, ovar ies.
6. Hor mones ar e r equir ed in specific amount . Their h yp os e cr e t i on (secr et ion in lesser
amount t han nor mal), as well as h yp e r s e cr e t i on (secr et ion in lar ger amount t han
nor mal) pr oduces ser ious physiological dist ur bances in t he body.
7. P r op e r t i e s of h or mon e s :
The hor mones have t he following pr oper t ies:
(a) They have low molecular weight .
(b) They ar e soluble in wat er and blood.
(c) They have no cumulat ive effect .
(d) They can act in ver y low concent r at ion.
(e) They ar e non-ant igenic.
(f) They ar e or ganic cat alyst s.
8. Ki n d s of h or mon e s : Based on t heir influence on physiological act ivit ies and cont r ol,
t he hor mones may be divided int o following 5 cat egor ies.
(i) Hor mones concer ned wit h met abolism e.g. Insulin et c.
(ii) Hor mones for gr owt h and development e.g. Somat ot r opin et c.
(iii) Hor omones of digest ion e.g. Gast r in, Secr et in et c.
(iv) Hor mones for r epr oduct ion e.g. Gonadot r opic and Sex hor mones.
(v) Hor mones t hat cont r ol ot her endocr ine glands e.g. Thyr ot r opin et c.
9. Bi och e mi ca l Cl a s s i fa ct i on of h or mon e s : All hor mones, depending upon t heir
chemical st r uct ur e, may be classified under t he following cat egor ies.
(i) Am i n e s : Hor mon es of pi n ea l gl a n d (mel a t on i n ) a n d a dr en a l medu l l a
(cat echolmines, viz., adr enaline, nonadr enaline).
(ii) Mod i fi e d Ami n o Aci d s : Hor mones of t hyr oid ar e iodinat ed t yr osine, e.g.,
t hyr oxine.
(iii) P e p t i d e s : Hor mones of hypot halamus (ARH, TRH, GRH, GIH), int er mediat e
(MSH) and post er ior lobes of pit uit ar y (ADH, oxyt ocin). ACTH of ant er ior pit uit ar y
a n d ca l ci t on i n of t h yr oi d bel on g t o t h i s ca t egor y. Th ey ma y fu r t h er be
differ ent iat ed int o shor t pept ides (e.g., oxyt ocin, ADH) and long pept ides (e.g.,
calcit onin, ACTH).
(iv) P r ot e i n s : Hor mones of pancr eas (e.g., insulin, glucagon), gast r oint est inal t r act ,
some female hor mones (e.g., r elaxin of ovar y and hCG of placent a), par at hor mone
(PTH) and most hor mones of ant er ior pit uit ar y except ACTH (e.g., TSH, FSH,
LA, LTH, GH) ar e pr ot einaceous.
(v) St e r oi d s : Hor mones a r e der ived fr om cholest er ol a nd ot her st er oids, e.g.,
aldost er one, cor t isol, sex cor t icoids (adr enal cor t ex), t est ost er one, est r adiol,
pr ogest er one (gonads except r elaxin, placent a except hCG).
10. RI A – Radioimmunoassay t echnique t o measur e t he hor mones, t heir pr ecur sor and
t heir met abolic end pr oduct s quant it at ively in a living body.
11. En d ocr i n e gl a n d s i n h u ma n b e i n gs : In man t he following major endocr ine glands
ar e pr esent .
(i) Pit uit ar y gland or Hypophysis
Physiology 151
(ii) Pineal Gland
(iii) Thyr oid gland
(iv) Par at hyr oid gland
(v) Adr enal gland
(vi) Gonads
(vii) Islet s of Langer hands in Pancr eas
(viii) Placent a
(ix) Thymus gland
(x) Hypot halamus
(xi) Gast r o-Int est inal lining
SUMMARY OF ENDOCRINE GLANDS AND THEIR HORMONES
I. Pituitary Gland
It consist s of t hr ee lobes, ant er ior , middle and post er ior .
Loca t i on : Floor of diencephalons and at t ached t o t he hypot halamus by a st alk t o for m
hypot halamic – pit uit ar y syst em.
(a ) An t e r i or l ob s of P i t u i t a r y: It possesses acidophils (a- cells), basophils (b-cells) and
chr omophobes (pr ecur sor s of ot her t wo). Acidophils pr oduce t wo hor mones (GH and
LTH) while basophils secr et e five hor mones (TSH, Lipot r opin, ACTH, FSH, LH).
(i) Soma t ot r op i n or Gr owt h St i mu l a t i n g Hor mon e (GH)
F u n ct i on s: St imulat es gr owt h by acceler at ing t he pr ot ein synt hesis and r et ent ion
of calcium.
Effe c t of h yp os e c r e t i on : Less secr et ion of GSH lea ds t o d w a r f i s m a nd
a cr omi cr i a in adult s.
Effe ct of Hyp e r se cr e t i on : 1. Gigant ism. Due t o excess secr et ion of GSH dur ing
childhood.
2. Acr omeagly in adult s.
(ii) Pr ol a ct i n / La ct ogen i c or Lu t eot r op h i c Hor m on e (LT H)– Pr ot eina ceous
hor mone (oft en included under gonadot r ophins) t hat st imulat es development of
mammar y glands dur ing pr egnancy and lact at ion aft er childbir t h. It is also called
mat er nit y hor mone.
(iii) Li pot r opi n or Ad i pok i n et i c Hor mon e– The hor mone st imulat es liber at ion of
fat t y acids fr om st or ed fat s.
(iv) Gon a d ot r opi n s: The hor mones, which r egulat e t he sexual act ivit y of gonads,
and ot her st r uct ur es of t he body. They include:
1. F ol l i cl e St i mu l a t i n g Hor mon e (F SH)
F u n ct i on s : St imulat ion of t est es in male t o for m t he sper ms fr om t he
semin ifer ou s t u bu les: I n fema les, t h e FSH st imu la t es ova r y for t h e
mat ur at ion of ovat ion follicle. Her e, it also st imulat es t he follicular cells t o
secr et e est r ogen . Th e la t t er is r espon sible for t h e secon da r y sexu a l
char act er s in female.
Effe ct of h yp os e cr e t i on : Failur e of gamet e for mat ion fr om t he gonads.
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2. I n t e s t i n a l Ce l l s St i mu l a t i n g Hor mon e (I CSH) i n ma l e s .
F u n ct i on s : St imulat es t he int er st it ial or Leydig’s cells or t est is t o pr oduce
t est ost er one. The lat t er br ings about secondar y sexual char act er s of male.
Effe ct of h yp os e cr e t i on : Impair ed development of t he ext er nal genit alia.
3. Le u t i n i zi n g h or mon e i n fe ma l e s (LH) (or Lu t r op i n ).
F u n ct i on s : Release of ovum fr om ovar y (ovulat ion); for mat ion of cor pus
lut eum and t he secr et ion of pr ogest er one fr om cor pus lut eum.
Effe ct of h yp os e cr e t i on : St er ilit y in females.
(v) Th yr oi d St i mu l a t i n g Hor mon e (TSH)
F u n ct i on : Pr oduct ion of t hyr oid hor mones fr om t hyr oid gland
(vi ) Ad r e n ocor t i ocot r op i n Hor mon e (ACTH):
F u n ct i on : Br ings about t he pr oduct ion and r elease of hor mones fr om adr enal
cor t ex.
(b) I n t e r me d i a t e l ob s of p i t u i t a r y
Hor mon e : Mel a n ocyt es st i m u l a t i n g h or m on e (MS H)
F u n ct i on : May be connect ed wit h pigment maint enance in hair , pr egnancy r elat ed
skin pigment at ion.
(c) P os t e r i or l ob s of p i t u i t a r y
Hormones
(i) Oxyt oci n (Pi t oci n ): St imulat ion of ut er ine cont r act ion dur ing child bir t h. St imulat ion
of milk flow by mammar y glands under t he cont r ol of Pr olact in or LTH.
(ii) Va sopr essi n (Pi t r essi n , ADH): Influences wat er balance by r educing out put of ur ine
by r eabsor pt ion of wat er fr om nephr ic filt r at e in DCH and CT. Also, causes ar t er iole
const r ict ion, r aise blood pr essur e and causes cont r act ion of sever al smoot h muscles
like t hose of int est ine, gall bladder , ur inar y bladder et c.
Hyper secretion
Causes Diabet es insipidus (Dr inker ’s disease) It is hor monal abnor malit y of p olyu r i a
(excessive ur ine) or mict ur at ing dilut e ur ine (hypot onic, sugar -fr ee) sever al t imes a
day.
II. Pineal Gland
It is a st alked small r ounded gland found behind t he ant er ior chor oid plexus on t he epit halamus.
Calcificat ion occur s in middle age. It has pineal and glial cells.
Hormones
The gland secr et es t wo biogenic amine hor mones
(i) S er ot on i n : Also by disint egr at ing blood cells. Const r ict s (Vasoconst r ict ion) blood vessel
at sit e of injur y and gives bur ning sensat ion.
(ii) Mel a t on i n :. The hor mone develops pale skin color in amphibians. It shows a mar ked
diur nal r hyt hm wit h maximum concent r at ion at midnight and minimum dur ing noon.
The hor mone cont r ols sleep, mood, ovar ian cycle, delays puber t y, opposes FSH and
LH hor mones (hence ant i-gonadot r ophic).
Physiology 153
III. Thyroid
Loca t i on : Base of lar ynx
Hor mon e s :
(i) Th yr oxi n e
(ii) T r i od ot h r on i n e (Bot h ar e iodinat ed amino acids)
F u n ct i on s : Regulat e cellular oxidat ion of food and basal met abolism, gr owt h and
differ ent iat ion.
Effe ct of h yp os e cr e t i on :
(i) Cr e t i n i s m in childr en, and Myxod e ma in adult .
(ii) Iodine deficiency goit er .
(iii) Ha s h i mot o Di s e a s e : It occur s in middle aged females due t o sensit izat ion of t heir
own t hyr oid pr ot ein called t hyr oglobulin. Ther e is mild hypot hyr oidism. The t hyr oid
gland is enlar ged.
Effe ct of Hyp e r se cr e t i on : Exop h t h a l a mi c goi t e r / Gr a ve d i s e a s e indicat ed by
pr ot r uding eyes, loss of weight , ner vous- ness and fast hear t beat .
(iv) Ca l ci t on i n (Pr ot ei n ): Regulat es calcium and phosphor us
IV. Parathyroids
Loca t i on : Behind t hyr oid
Hor mon e s : Par at hor mone (pr ot ein)
F u n ct i on s : Regulat es calcium and phosphor us balance in t he blood.
Effe ct of h yp os e cr e t i on : Reduces Ca in blood and r esult s in sever e cr amps Par at hyr oid
t et any.
Effe ct of Hyp e r se cr e t i on : High Ca level in blood by r eleasing it fr om bones and leading
t o ost est is fibr osa cyst ica.
V. Adrenals
Each adr enal has t wo par t s, an out er cor t ex and inner medulla.
Loca t i on : Over t he super ior end of kidneys.
(a ) Ad r e n a l Cor t e x. The out er par t of adr enals which fur t her consist s of t hr ee zones
(i ) Zon a gl ome r u l os a :
Hor mon e s : Mi n er a l ocor t i coi d s (m a i n l y a l d ost er on e)
F u n ct i on s : Regulat ion of Na
+
and K
+
in blood. Incr eases Na
+
level, and lower s
t hat of K
+
(i i ) Zon a F a s ci cu l a t a
Hor mon e : Gl u cocor t i coi d s (Ma i n l y cor t i sol )
F u n c t i on : Incr ease glucose level in body by changing pr ot eins and fast t o
car bohydr at es.
Hyp e r s e cr e t i on : Cushing’s syndr ome due t o high glucose level, and high Na
+
level caused by excess of glucocor t icoids. High B.P. and r ise in blood volume.
(i i i ) Zon a r e t i cu l os a
Hor mon e : An d r ogen s (Ma l e sex h or m on es)
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F u n ct i on : Development of male secondar y sexual char act er s.
Hyp e r s e cr e t i on : Adr enal vir ilism due t o excess of andr ogens secr et ed in female
who develops cer t ain masculine feat ur es.
(b) Ad r e n a l me d u l l a (The inner por t ion of adr enals)
The adr enals ar e commonly called t he glands of emer gency because of t he r ole of
t heir hor mones.
Hor mon e:
(i) Nor a d r en a l i n e (= Nor ep i n ep h r i n e) Secr et ed under nor mal condit ions.
F u n ct i on : Regulat e blood pr essur e under nor mal condit ion by const r ict ing small
ar t er ies.
(ii) Ad r en a l i n e (= Ep i n ep h r i n e) secr et ed under condit ions of physical and ment al
st r ess such as gr ief, pain, injur y, anger , fall in blood pr essur e and blood sugar ,
cold fea r , et c. I n ma n, a dr ena lin is pr oduced in much mor e a mount t ha n
nonadr enaline.
F u n ct i on : Incr ease of hear t beat r at e, br eat hing r at e and blood sugar . Dilat ion
of small ar t er ies of hear t , br ain and skelet al muscles. Const r ict ion of small
ar t er ies of skin: const r ict ion of small ar t er ies of skin; cont r act ion of spleen t o
squeeze out t he r eser ve, dilat ion of pupil, cont r act ion of ar r ect or pili muscles
(goose flesh condit ion).
VI. Gonads
(a ) Te s t e s . A het er ocr ine gland. The endocr ine por t ion is r epr esent ed by t he Leydig’s
cells (Int er st it ial cells) pr esent bet ween t he seminifer ous t ubules.
Loca t i on : Sit uat ed in Scr ot um
Hor mon e s : T est ost er on e a n d ot h er a n d r ogen s.
F u n ct i on s :
1. Gr owt h of t he r epr oduct ive syst em t o full size and t o funct ional st at e.
2. St imulat es sper m for mat ion.
3. For mat ion of sexual char act er s of female.
Hyp os e c r e t i on : Eunuchoidism (I na dequa t e for ma t ion of r epr oduct ive syst em,
incapabilit y of sper m for mat ion).
(b) Ova r i es
Loca t i on : In t he abdominal cavit y.
Hor mon e s :
(i ) Oe s t r oge n fr om fol l i cu l a r ce l l s .
F u n ct i on : Gr owt h of fema le r epr oduct ive syst em t o full size, for ma t ion of
secondar y sexual char act er s of female.
(i i ) P r oge s t e r on e (s e cr e t e d b y cor p u s l u t e u m).
F u n ct i on :
(a) Suspends t he ovulat ion.
(b) For mat ion of placent a.
(i i i ) Rel a xi n (pr oduced by cor pus lut eum at t he end of pr egnancy)
Physiology 155
F u n ct i on : Cont r ols t he development of foet us, Relaxat ion of cer vix, and ligament s
of pelvic gir dle t o facilit at e t he bir t h.
VII. Pancreas
A myxocr ine gland. It s endocr ine par t is r epr esent ed by Isle t s of Langer hans which have t wo
t ype of cells t he a (alpha) cells, and b (bet a) cells.
Loca t i on : In t he duodenal loop.
(a ) Be t a ce lls
Hor mon e : I n su l i n (an anabolic hor mone)
F u n ct i on :
(i) Conver sion of glucose t o glycogen in liver and muscles.
(ii) Pr omot es pr ot ein synt hesis in t issues.
(iii) Pr omot es synt hesis of fat s fr om fat t y acids
(iv) Inhibit s br eakdown of pr ot eins and fat s.
Hyp os e cr e t i on : Diabet es mellit us indicat ed by t he r ise of glucose level in blood, and
higher level of cholest er ol, excr et ion of sugar by t he kidneys in ur ine; excessive
t hir st ; delayed wound healing of pr ot eins in t issues, and gangr ene.
(b) Al p h a ce l l s
Hor mon e : Glucagon
F u n ct i on :
(i) Conver t s glycogen t o blood glucose in liver
(ii) For mat ion of glucose fr om amino acids.
VIII. Placenta
Loca t i on : A living connect ion bet ween t he foet us and ut er ine wall.
Hor mon e s :
(i ) Hu ma n ch or i on i c gon a d ot r op i n (HCG)
F u n ct i on : Pr oduced in lar ge amount in fir st few mont hs of pr egnancy. It st imulat es
t he enlar gement of cor pus lut eum t o secr et e pr ogest er one. Pr esence of hCG in ur ine
is indicat ion of pr egnancy, used for pr egnancy t est
(ii) Pr ogest er on e: Funct ionally same as t he one pr oduced by cor pus lut eum.
(iii) Est r ogen : : Similar r ole as t hat pr oduced by t he follicular cells of ovar y
IX. Thymus
Loca t i on : Near t he hear t . Get s r educed in adult s.
Hor mon e : Th ymosi n (Pr ot ei n )
F u n ct i on : Pr olifer at ion of lymphocyt es, and also r egulat es gr owt h; impor t ant for immune
syst em.
Hyposecr et ion: Poor development of immunit y. Ret ar ded development of sex or gans.
X. Hypothalamus
(Composed of neur osecr et or y cells of t he floor of diencephalons secr et e neur ohor mones).
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Loca t i on : Floor of diencephalons of br ain; connect ed t o t he ant er ior lobe of pit uit ar y by
hypophyseal blood vessel, and wit h post er ior lobe of pit uit ar y t hr ough t he axons of it s neur ons.
Some hor mones ar e pour ed int o adenohypophysis (ant er ior , and int er mediat e pit uit ar y), t hr ough
hypophysial por t al syst em while t wo hor mones (oxyt oci n a n d ADH) ar e dir ect ly t aken by
ner ve cells in neur ohypophysis (post er ior pit uit ar y). Hor mones pour ed in adenohypophysis
funct ion as r eleasing or inhibit ing hor mones. They ar e pept ide in nat ur e.
Hor mon e s :
(i ) Th yr ot r opi n Rel ea si n g Hor m on e (T-RH): Release of TSH fr om ant er ior pit uit ar y.
(i i ) Ad r en ocor t i cot r opi n - Rel ea si n g Hor m on e (A-RH): Release of ACTH fr om ant er ior
pit uit ar y.
(i i i ) S om a t ot r opi n R el ea si n g h or m on e (S -R H): Release of SH fr om ant er ior pit uit ar y.
(i v) Gon a d ot r opi n - R el ea si n g h or m on e (G-R H): Release of goandot r opins fr om t he
ant er ior pit uit ar y.
(v) Gr owt h i n h i bi t i n g h or m on e (GI H) or som a t ost a t i n : Inhibit s t he hor mone fr om
pit uit ar y.
XI. Gastrointestinal lining
Loca t i on : Inner layer of wall of gut .
(a ) St oma ch
Hor mon e : Ga st r i n
F u n ct i on : St imulat ion of gast r ic glands t o r elease gast r ic juice. Also st imulat es
st oma ch movement s.
(b) I n t e s t i n e
(i ) S ecr et i n
(1) St imulat es pancr eas t o r elease enzymes.
(2) St imulat es gall bladder t o r elease bile.
(i i ) Ch ol ecyst ok i n i n Pa n cr eoz ym i n (CCK – PZ): St ops t he secr et ion of gast r ic
juice and slows down gast r ic movement s.
(i i i ) En t er oga st r on e: St ops t he secr et ion of gast r ic juice and slows down gast r ic
movement s.
(i v) En t er ocr i n i n n : Release t he int est inal juice fr om cr ypt s of Liber kuhn.
(v) Du ocr i n i n : Release of mucus fr om Br unner ’s Glands
(vi ) Vi l l i k r i n i n : Acceler at es t he movement of villi.
Local Hormones (Tissue Hormones)
(i ) Local hor mones ar e t hose hor mones which do not pass int o blood but diffuse t o t he
t ar get as t he same is near by. They ar e also called par acr ines or par ahor mones
(i i ) The impor t ant ones ar e acet ylcholine (fr om ner ve endings), kinins (br adykinins) and
pr ost aglandins. Pr ost aglandins ar e fat t y acid der ivat ives, which have diver se funct ions
including blood flow r egulat ion, ut er ine cont r act ions, ant it hr ombot ic, r elaxat ion of
ar t er ial smoot h muscles, et c. Kinins or br adykinins ar e polypept ides t hat cause
cont r act ion of smoot h muscles and dilat ion of blood vessels.
(i i i ) The gaseous hor mone, nit r ic oxide (also an air pollut ant ), is also a local hor mone.
Molecule of t he year 1994, it has been used in r escuing end st age pat ient s in r espir at or y
Physiology 157
dist r ess. It is a signalling molecule of car diovascular syst em t hat r egulat es blood
pr essur e and blood flow besides r elaxing smoot h muscle cells. It is cur r ent ly used in
male pot ency dr ug viagr a.
Pheromones
(i ) They ar e chemicals used for communicat ion amongst individuals of t he same species.
Pher omones ar e pr oduced by exocr ine glands and ar e t r ansmit t ed t o ot her member s
t hr ough air or excr ement s (e.g., ur ine in dogs).
(i i ) Pher omones invoke a specific r esponse in ot her member s like r ecognit ion, at t r act ion,
war ning, et c. Bombykol is sex at t r act ant pher omone pr oduced by female silkwor m
mot h, which is r eceived by male ant enna sever al kilomet er s away.
(i i i ) Ant t r ail is due t o pher omone. Queen Bee is r ecognized by wor ker s due t o r elease of
9-oxydecenoic acid by it . Dor mit or y effect is also believed t o be due t o pher omones.
REPRODUCTION
Repr oduct ion is t he for mat ion of new similar young living or ganisms by t he gr own up individuals
of a species or r ace. It is meant for per pet uat ion of t he r ace/species because individuals ar e
bound t o die aft er a life span. Repr oduct ion pr ovides gr oup immor t alit y. Four pr ocesses ar e
basic t o r epr oduct ion – DNA r eplicat ion, cell division, for mat ion of r epr oduct ive unit s and
development of a new individual. Animal r epr oduct ion is of t wo t ypes, asexual and sexual.
Asexual Reproduction
It is a mode of r epr oduct ion or for mat ion of new young individuals fr om a specialized or
unspecialized par t of single par ent wit hout t he for mat ion and fusion of gamet es. Besides being
unipar ent al and absence of gamet es, asexual r epr oduct ion is char act er ized by divisions t hr ough
mit osis only, genet ic similar it y bet ween par ent and young ones, r apidit y and absence of haploid-
diploid alt er at ion. Asexual r epr oduct ive pr opagule is called blast os. It cont ains t ot ipot ent cells
like a r ch a e ocyt e s (sponges), i n t e r s t i t i a l ce l l s (cnidar ia), p a r e n ch yma (plat yhelmint hes)
and n e ob l a s t s (annelids), et c. An individual pr oduced t hr ough asexual r epr oduct ion is r amet .
Clone is gr oup of all genet ically similar individuals for med t hr ough asexual r epr oduct ion.
1. F i s s i on : It is a t ype of asexual r epr oduct ion in which t he body of an individual
under goes division t o pr oduce t wo or mor e equal sized daught er s.
(i ) Bi n a r y Fi ssi on . The mat ur e individual divides int o t wo equal sized daught er
individuals. Binar y fission is ir r egular (can occur in any plane) in Amoeba,
longit udinal in Euglena, oblique in dinoflagellat es and t r ansver se in Paramecium
and Planaria. In unicellular for ms, binar y fission is accomplished t hr ough mit ot ic
nuclear division followed by cyt okinesis. In mult icellular individuals like Planaria
t he post er ior par t is fixed fir mly t o subst r at um while ant er ior par t ext ends for war d
and exer t s a pull causing t he middle par t t o br eak.
(i i ) Mu l t i pl e Fi ssi on : It is for mat ion of a number of small daught er s by division of
a par ent . In unicellular for ms, t he nucleus divides a few t imes followed by
collect ion of cyt oplasm ar ound each daught er nucleus for ming a number of
daught er cells, e.g., Amoeba, Plasmodium, Monocyst is.
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(i i i ) Pl a sm ot om y: Cleavage of plasmodium or mult inucleat e body int o mult inucleat e
par t s, e.g., Opalina, Pelomyxa.
(i v) S t r obi l i sa t i on : A modificat ion of mult iple fission is st r obilisat ion in mult icellular
animals (body segment at ion for r epr oduct ion as for ming zooids in coelent er at es
like Aurelia and pr oglot t ides in Tapewor m).
2. Cys t s a n d Sp or e s : They ar e minut e pr opagules, which funct ion as dissemules as
well as per ennat ing st r uct ur es. Each cyst or spor e gives r ise t o a new individual.
3. Bu d d i n g: It is a mode of asexual r epr oduct ion in which new or ganisms develop as
out gr owt h or buds fr om a par ent .
(i ) Ex ogen ou s/ Ex t er n a l Bu d d i n g. The bud develops ext er nally on t he par ent . In
t he young condit ion t he t wo ar e int er nally connect ed t o pr ovide nour ishment t o
t he bud. Lat er on, t he bud enlar ges and becomes nut r it ionally independent . It
t hen separ at es as a new individual, e.g., Yeast , Hydra. However , in some cases,
t he new individual r emains joined t o t he par ent t o for m colony, e.g., S cypha.
(i i ) En d ogen ou s/ I n t er n a l Bu d d i n g. Int er nal buds or gemmules/ st at oblast s occur
in sponges/br yozoans t o help in disper sal as well as per ennat ion.
4. F r a gme n t a t i on : It is br eaking of an individual int o t wo or mor e par t s due t o ext er nal
(e.g., St ar fish) or int er nal for ces (e.g., Tunicat es). Each fr agment gr ows int o a new
individual.
Sexual Reproduction
It is mode of mult iplicat ion in which t he new individuals or young ones ar e for med t hr ough t he
pr ocess of for mat ion and fusion of gamet es. Haploid (gamet es)- diploid (individual) alt er nat ion
occur s. Ga met ogenesis involves meiosis. Fusion of ga met es or fer t iliza t ion r est or es t he
chr omosome number . The offspr ing is pr oduced by t he gr owt h of t he fusion pr oduct of gamet es
called zygot e. Sexual r epr oduct ion is commonly bipar ent al. A number or var iat ions appear in
t he offspr ing.
Parthenogenesis (Apomixis)
It is t he development of a new individual fr om a single gamet e (gener ally t he egg/ovum) wit hout
involving fer t ilizat ion. On t he basis of chr omosome set s, par t henogenesis is of t wo t ypes:
(i ) Ar r h en ot ok y (Haploid Par t henogenesis). Haploid eggs gr ow t o for m haploid males,
e.g., ar achnids, some insect s.
(i i ) Th el ot ok y (Diploid Par t henogenesis). Diploid eggs gr ow wit hout fer t ilizat ion int o
diploid individuals, gener ally females, e.g., Gall Fly
Par t henogenesis can be nat ur al or ar t ificial. Nat ur al par t henogenesis may be obligat or y
or cyclic.
Natural Parthenogenesis
1. Ob l i ga t or y/Comp l e t e P a r t h e n oge n e s i s : Ma les a r e a bsent . Fema les develop
par t henogenet ically, e.g. r ot ifer s, Typhlina brahmina (small lizar d), Lacerta saxicola-
armeniaca (Caucasian Rock Lizar d), Cnemidophorus (Whipt ail lizar d of Amer ica)
2. Cycli c/I n comp le t e P a r t h e n oge n e s i s : Bot h sexual and par t henogenet ic individuals
occur . In aphids, sever al gener at ions of par t henogenet ic females develop followed by
Physiology 159
for mat ion of bot h males and females t o per for m sexual r epr oduct ion. In Tur key, 40%
of t he males develop par t henogenet ically. In Honey Bee, male or dr one develops
par t henogenet ically (no meiosis at t he t ime of sper mat ogenesis) while queen and
wor ker s develop fr om fer t ilized eggs. Also in wasps and ant s. In Gall Fly, lar vae may
lay eggs t hat develop par t henogenet ically (paedogenesis).
Artificial/Induced Parthenogenesis
Sugar , salt , alkaloids and ot her chemicals, heat , cold, pr icking and ot her st imuli can st imulat e
eggs t o under go cleavage and for m par t henogenet ic embr yos. Being haploid, t hey gener ally do
not sur vive.
GAMETOGENESIS
The pr ocess of for mat ion off gamet es in t he gonads is called gamet ogenesis. Gamet es ar e
pr oduced by way of meiot ic division r esult ing in t he r educt ion of number of chr omosomes t o
one half. The pr oduct ion of sper ms in t he seminifer ous t ubules of t est is is called sper mat ogenesis
and pr oduct ion of ova in ovar ies is called oogenesis.
1. Sp e r ma t oge n e s i s
Sper mat ogenesis is t he pr ocess of for mat ion of haploid funct ional sper mat ozoa fr om
diploid ger minal cells of seminifer ous t ubules. Lining layer of seminifer ous t ubules
possesses pr imar y ger m cells and indiffer ent cells t hat mat ur e int o nur se cells or
Ser t oli cells.
(i ) Mu l t i p l i ca t i on Ph a se: Diploid pr imar y ger m cells under go r epeat ed mit osis t o
for m a number of diploid sper mat ogonia.
(i i ) S p er m a t ocyt ogen esi s: Each sper mat ogonia divided mit ot ically t o fr om t wo
int er connect ed pr imar y sper mat ocyt e
(i i i ) Gr owt h Ph a se: Pr imar y sper mat ocyt es gr ow in size.
(i v) Ma t u r a t i on Ph a se: Each diploid pr imar y sper mat ocyt e under goes meiosis I t o
for m t wo haploid secondar y sper mat ocyt es. All t he secondar y sper mat ocyt es
der ived fr om a single sper mat ogonium r emain at t ached t o one anot her . Secondar y
sper mat ocyt es- divides by meiosis II, each giving r ise t o t wo haploid sper mat ids.
The sper mat ids become par t ially embedded in Ser t oli cells for nour ishment and
suppor t .
(v) S p er m i ogen esi s: Sper miogenesis is differ ent iat ion of a sper mat ozoon fr om a
sper mat id. Golgi appar at us for ms acr osome. Cent r iole divides int o t wo. Dist al
cent r iole for ms axial filament . Mit ochondr ia collect ar ound upper par t of axial
filament . Nucleus under goes condensat ion. A sper mat ozoon now separ at es while
t he unused cyt oplasm degener at es. Heads of sper mat ozoa r emain embedded for
some t ime in Ser t oli cells but ult imat ely t he sper mat ozoa ar e r eleased int o
lumen of seminifer ous t ubule for onwar d passage.
Sperm
Human sper m is dar t -like flagellat e st r uct ur e of 60 µm lengt h. It has four par t s—
(i ) Hea d . Knob-like t er minal par t has t wo component s, acr osome and nucleus. Acr osome
cont ains sper m lysins. Nucleus is compact mass of DNA having some p r ot a mi n e s .
On t he out side is pr esent a double membr ane head cap.
160 CSIR-NET Life Sciences
(i i ) Neck . It is shor t nar r ow par t bet ween head and middle piece, which cont ains t wo
cent r ioles, unconnect ed pr oximal cent r iole and dist al cent r iole at t ached t o axial
filament (t hat passes int o middle piece).
(i i i ) Mi d d le P i ece. It is cylindr ical par t . It has axial filament sur r ounded by 10-14 spir al
t ur ns of mit ochondr ia and bear ing t owar ds t he end a r ing cent r iole or annulus.
Mit ochondr ia pr ovide ener gy for swimming but food is limit ed.
(i v) Ta i l . It is nar r ow vibr at ile long par t , wit h t wo r egions, main and end piece. Main
piece of t ail is 0.5 mm in diamet er near t he beginning but gr adually nar r ows behind.
It has an axial filament , small amount of cyt oplasm and plasma membr ane. In t he
end piece, cyt oplasm and membr ane ar e absent .
2. Oogenesis
It is t he pr ocess of for mat ion, development and mat ur at ion of haploid ova fr om diploid ger minal
cells of ovar y. Oogenesis occur s in t hr ee phases
(i ) Mu l t i p l i ca t i on P h a s e . Diploid pr imar y ger m cells fr om ger minal epit helium of
ovar y mult iply mit ot ically and for m oogonia. The lat t er pr oduce oviger ous cor ds or
egg t ubes of Pfluger in mammals.
(i i ) Gr owt h P h a s e . It is pr olonged and slow. Oogonia for m r ounded masses or egg nest s
at t he t ips of egg t ubes of Pfluger . An egg nest for ms ovar ian follicle. One cent r al
oogonium gr ows and funct ions as pr imar y oocyt e. The ot her s for m t he cover ing
follicular cells. The lat t er pr ovide nour ishment t o pr imar y oocyt e. Some nour ishment
also comes fr om out side. Enlar ged pr imar y oocyt e secr et es mucopr ot ein membr ane
or zona pellucida out side it s own plasma membr ane or vit elline membr ane. Ther e is
incr ease in r eser ve food, size of nucleus, number of mit ochondr ia, funct ioning of
Golgi appar at us and complexit y of endoplasmic r et iculum.
(i i i ) Ma t u r a t i on P h a se. Meiosis occur s. Nucleus shift s t owar ds animal pole and under goes
meiosis I. A daught er nucleus alongwit h small quant it y of cyt oplasm is ext r uded as
pr imar y polar body or polocyt e below zona pellucida. Simult aneously pr imar y oocyt e
is changed int o haploid secondar y oocyt e. Ovum is gener ally shed in secondar y oocyt e
st age. Meiosis II is complet ed in fallopian t ube at t he t ime of fer t ilizat ion. It t hen
pr oduces a small secondar y polar body and a lar ge ovum. The pr imar y polar body
may also divide.
Ovum
Human egg or ovum is noncleidoic and alecit hal (near ly micr olecit hal), r ounded female gamet e
having a diamet er of about 100 mm. The ovum possesses t hr ee cover ings – inner t hickened
plasma membr ane or vit elline membr ane, middle mucopr ot ein zona pellucida and out er cellular
cor ona r adiat a wit h r adially elongat ed scat t er ed cells held in mucopolysacchar ide (hyalur onic
acid). In bet ween plasma membr ane ( = vit elline membr ane) and zona pellucida is per i-vit elline
space in which ar e pr esent 1- 3 polar bodies t owar ds animal pole. The opposit e end is veget al
pole. Cyt oplasm of ovum is called ooplasm. It has a lar ge nucleus or ger minal vesicle. Ect oplasm
possesses
Physiology 161
FERTILIZATION IN HUMANS
Only a single ovum is r eleased in human females fr om one of t he t wo ovar ies t owar ds t he
middle of ovar ian/menst r ual cycle. It passes int o fallopian t ube and r est s inside ampulla for
some t ime. Human male pr oduces 300 - 400 million sper ms per ejaculat ion. They ar e deposit ed
in vagina dur ing coit us. The pr ocess of deposit ion of sper ms in t he female genit al t r act is called
inseminat ion. A number of t hem ar e demobilised or eat en.
Fer t ilizat ion involves t he following st eps:
(i ) Ap p r oxi ma t i on of Sp e r m a n d Ovu m. Sper ms can r emain mot ile for 24-48 hour s.
They swim at t he r at e of 1. 5 - 3.0 mm/min. They ar e able t o r each t he ampulla par t
of female genit al t r act par t ly by t heir own swimming and par t ly by cont r act ion of
ut er us and fallopian t ubes. One sper m comes t o lie against t he ovum and under go
fer t ilizin (fr om ovum) and ant ifer t ilizin (fr om sper m) compat ibilit y r eact ion (Lillie,
1919) in t he r egion of animal pole.
(i i ) Acr os ome Re a ct i on . It pr epar es t he sper m t o fer t ilize ovum. The phenomenon is
called capacit at ion. In cont act wit h cor ona r adiat a, t he acr osome cover ing lyses t o
r elease acr osome. Wit h t he help of sper m lysins (chiefly hyalur onidase), acr osome
dissolves cor ona r adiat a and zona pellucida in fr ont of it .
(i i i ) Egg Re a ct i on . A small pr ot uber ance or fer t ilizat ion cone (cone of r ecept ion), develops
fr om t he sur face of ovum in t he r egion of animal pole.
(i v) P e n e t r a t i on of Sp e r m. Sper m head est ablishes cont act wit h lat er al sur face of
fer t ilizat ion cone. Plasma membr anes of t he t wo dissolve. Cont ent s of head (nucleus),
neck and middle piece of sper m ent er ooplasm. Tail is left out side. Fer t ilizat ion cone
subsides. Depolar isat ion of egg membr ane kills ot her sper ms. Plasma membr ane of
t he egg is now modilled wit h t he help of mucopolysacchar ide cor t ical gr anules int o
fer t ilizat ion membr ane. A per ivit elline space is cr eat ed bet ween it and zona pellucida.
(v) Act i va t i on of Ovu m. Ovum (pr eviously in secondar y oocyt e st age) under goes meiosis
II and ext r udes a secondar y polar body. It is now t he act ual ovum or female gamet e.
(vi ) F u s i on of Sp e r m a n d Egg Nu cl e i . The envelopes of t he sper m and egg nuclei
degener at e and t heir chr omosomes int er mingle t o for m ‘synkar yon’. The act is called
kar yogamy or syngamy. The pr oximal cent r iole br ought by sper m help~ for m t he
spindle for t he division of synkar yon (cleavage nucleus). Fer t ilized egg is also called
zygot e
Practice Test Paper-I
1. Miner al nut r ient s
(a) ar e not essent ial t o plant gr owt h, since all a plant needs is wat er and CO
2
.
(b) cont r ibut e lit t le t o t he weight of a plant .
(c) ent er plant s via t he st omat a.
(d) ar e or ganic nut r ient s.
2. Pr ofessor Ar un claims t o have discover ed a new macr onut r ient r equir ed for plant
gr owt h. Most of Pr ofessor Ar un’s colleagues ar e skept ical of t his claim. Why might
t hey consider it unlikely?
(a) All t he nut r ient s r equir ed for plant gr owt h have alr eady been found.
(b) It is ver y difficult t o pr ove t hat a plant needs a cer t ain nut r ient .
(c) Plant s need t housands of nut r ient s; a new one is not significant .
(d) Any nut r ient needed in lar ge amount s has pr obably been found alr eady.
3. Soil could be deficient in any of t he following nut r ient s. If you had t o supply one of
t hem, which would be needed in t he smallest amount ?
(a) ir on (b) phosphor us
(c) nit r ogen (d) pot assium
4. Which of t he following is a sympt om of magnesium deficiency?
(a) yellowing of younger leaves’ pr ior t o yellowing of older leaves.
(b) enhanced plant gr owt h, since magnesium is t oxic t o plant s.
(c) chlor osis
(d) decr eased t r anspir at ion
5. Topsoil
(a) is t he r elat ively iner t upper layer of soil.
(b) does not r et ain wat er .
(c) is a mixt ur e of r ock fr agment s, living or ganisms, and humus.
(d) is unifor m in t ext ur e.
6. The r oot s of many aquat ic plant s have special st r uct ur es t hat pr oject above t he sur face
of t he wat er . For example, cypr ess t r ees (which gr ow in swamps) have knees t hat
ext end upwar d above wat er level. Which of t he following is t he most logical funct ion
of t hese st r uct ur es?
(a) obt aining car bon dioxide for phot osynt hesis
(b) nit r ogen fixat ion
(c) obt aining oxygen for t he r oot s
(d) t r anspir at ion
7. The clay par t icles in soil ar e impor t ant because t hey
(a) ar e composed of nit r ogen needed by plant s.
(b) eliminat e spaces for air and facilit at e dr ainage.
Practice Test Paper–Physiology 163
(c) fill spaces and keep oxygen out of t he soil.
(d) ar e char ged and hold ions needed by plant s.
8. Soil can easily become deficient in _____, because t hese ions ar e negat ively char ged
and do not st ick t o negat ively char ged clay par t icles.
(a) pot assium. (b) calcium.
(c) magnesium. (d) nit r at e.
9. Fer t ilizer s ar e usually enr iched in
(a) ir on, manganese, and zinc.
(b) calcium, bor on, and car bon.
(c) nit r ogen, phosphor us, and pot assium.
(d) molybdenum, copper , and magnesium.
10. Nit r ogen-fixing bact er ia in t he soil
(a) conver t nit r at es t o N
2
.
(b) conver t ammonia int o ammonium.
(c) conver t at mospher ic nit r ogen int o ammonia.
(d) use nit r at es t o make amino acids t hat plant s can use.
11. Nit r ogen fixat ion is
(a) using nit r ogen t o build molecules such as pr ot eins and nucleic acids.
(b) conver t ing nit r ogen in t he air int o a for m usable by plant s.
(c) r ecycling nit r ogen fr om or ganic mat t er in t he soil.
(d) absor bing N
2
fr om t he soil.
12. The enzyme t hat cat alyzes t he conver sion of at mospher ic nit r ogen int o ammonia is
(a) cat alase. (b) nit r ogenase.
(c) r educt ase. (d) r ubisco.
13. The r elat ionship bet ween legumes and Rhizobium is
(a) mut ualist ic. (b) par asit ic.
(c) compet it ive. (d) commensalism
14. Mycor r hizae ar e
(a) nut r ient s r equir ed by plant s in r elat ively small amount s.
(b) plant s such as mist let oe t hat par asit ize ot her plant s.
(c) cells t hat cont r ol evapor at ion of wat er fr om leaves.
(d) associat ions of r oot s wit h beneficial fungi.
15. The t er m alt er nat ion of gener at ions r efer s t o a plant ’s life cycle alt er nat ing bet ween
(a) t he pr oduct ion of haploid gamet es by meiosis wit h t he pr oduct ion of diploid spor es
by mit osis.
(b) a haploid gamet ophyt e gener at ion and a haploid spor ophyt e gener at ion.
(c) a haploid gamet ophyt e gener at ion and a diploid spor ophyt e gener at ion.
(d) a flower pr oducing gener at ion and a leaf-pr oducing gener at ion.
164 CSIR-NET Life Sciences
16. Self-incompat ibilit y
(a) wor ks t he same way in all plant s.
(b) ar e all based on t he same mechanism of t r ansplant r eject ion seen in animals.
(c) maint ains var iat ion.
(d) is t he r eject ion of a gr aft by a plant .
17. In angiosper ms, each pollen gr ain pr oduces t wo sper m. What do t hese sper m do?
(a) Each one fer t ilizes a separ at e egg cell.
(b) One fer t ilizes an egg and t he ot her fer t ilizes t he fr uit .
(c) One fer t ilizes an egg and t he ot her is kept in r eser ve.
(d) One fer t ilizes an egg and t he ot her fer t ilizes a cell t hat develops int o st or ed food.
18. What is endosper m?
(a) male r epr oduct ive cells in plant s
(b) st or ed food in a seed
(c) cells t hat make up t he bulk of a pollen gr ain
(d) t he fleshy par t of a fr uit such as an apple or st r awber r y
19. The cor r ect ar r angement of t he mer ist emat ic t issue of an embr yo is
(a) a r ing of lat er al mer ist em sur r ounding apical mer ist em.
(b) a r ing of pr ocambium sur r ounding a r ing of mer ist em sur r ounding a r ing of
pr ot oder m.
(c) a r ing of hypocot yl sur r ounding a r ing of r adicle sur r ounding a r ing of epicot yl.
(d) a r ing of pr ot oder m sur r ounding a r ing of gr ound mer ist em sur r ounding
pr ocambium.
20. The scut ellum
(a) is a specialized cot yledon found in cer t ain monocot s.
(b) develops int o t he seed coat .
(c) pr esent s a bar r ier t o self-fer t ilizat ion.
(d) is a specialized cot yledon found in dicot s.
21. Aft er fer t ilizat ion, t he _____ develops int o a seed and t he _____ develops int o a fr uit .
(a) ovule . . . ova r y
(b) pollen gr ain . . . ovule
(c) ovar y . . . ovule
(d) egg . . . ovule
22. Plant s gr owing in har sh envir onment s such as deser t s, sand dunes, and ar ct ic t undr a
oft en r epr oduce veget at ively. This is because
(a) t her e ar e few animals available t o pollinat e t hem.
(b) t hey ar e member s of plant families t hat only r epr oduce asexually.
(c) fr uit s would fr eeze or dr y out in t hese envir onment s.
(d) veget at ive r epr oduct ion is not as r isky as making seeds.
Practice Test Paper–Physiology 165
23. In cont r ast t o animals, plant s exhibit ______ and _____.
(a) det er minat e gr owt h . . . per sist ent mor phogenesis
(b) indet er minat e gr owt h . . . ar e not capable of mor phogenesis t hr oughout t heir
lifespan
(c) indet er minat e gr owt h . . . per sist ent mor phogenesis
(d) gr owt h . . . mor phogenesis
24. Which of t he following seedlings will pr obably bend t owar d t he light ?
(a) t ip cover ed wit h a cap made of black plast ic
(b) t ip separ at ed fr om base by a gelat in block
(c) t ip separ at ed fr om base by aluminum foil
(d) t ip cut off and place a block of agar over half of t he cut por t ion; t he side wit h t he
agar block will bend t owar ds t he light
25. In shoot s, br anching is inhibit ed by _____ fr om t he t ip of a gr owing shoot , but t his
effect is count er ed by _____ fr om t he r oot s.
(a) cyt okinins . . . auxins
(b) gibber ellins . . . et hylene
(c) auxins . . . cyt okinins
(d) gibber ellins . . . abscisic acid
26. As leaf let t uce mat ur es, t he basal edible leaves suddenly send up a t all flower ing
shoot . Aft er t he plant bolt s like t his, it no longer pr oduces br oad, t ast y leaves. Suppose
you want ed t o pr event bolt ing so t hat you could har vest let t uce longer . You might
look for some way t o int er fer e wit h t he effect s of
(a) abscisic acid. (b) gibber ellins.
(c) cyt okinins. (d) et hylene.
27. Seeds of many deser t plant s will not ger minat e unt il a heavy r ain washes away t heir
(a) phyt ochr ome. (b) abscisic acid.
(c) gibber ellins. (d) auxins.
28. The abscission layer
(a) causes a shoot t o bend t owar d light .
(b) secr et es cyt okinin.
(c) is t he locat ion of t he biological clock in a plant .
(d) is wher e a leaf separ at es fr om a st em.
29. When a plant st r uct ur e such as a leaf is injur ed, it pr oduces ______, which may cause
t he par t t o age and dr op off.
(a) cyt okinins (b) et hylene
(c) auxins (d) abscisic acid
30. Once a flower is pollinat ed, changes occur t hat make it less at t r act ive t o insect s. It s
pet als, for example, shr ivel and fall off. Pollinat ion must
(a) incr ease t he out put of cyt okinins in t he flower .
(b) block t he flow of auxins fr om t he r oot s.
166 CSIR-NET Life Sciences
(c) t r igger t he r elease of et hylene in t he flower .
(d) incr ease t he for mat ion of phyt ochr ome, which set s t he biological clock.
31. In t he aut umn, t he amount of ____ incr eases and t he amount of ____ decr eases in
fr uit and leaf st alks, causing a plant t o dr op fr uit and leaves.
(a) et hylene . . . auxin
(b) gibber ellin . . . abscisic acid
(c) cyt okinin . . . abscisic acid
(d) auxin . . . et hylene
32. Plant hor mones act by affect ing t he act ivit ies of
(a) genes. (b) genes and enzymes.
(c) enzymes. (d) genes, membr anes, and enzymes.
33. _______ appear t o be r esponsible for gr avit r opism.
(a) St at olit hs (b) phyt ochr omes
(c) Gibber ellins (d) Cyt okinins
34. A r apid loss of wat er in specialized cells in t he sensit ive plant Mimosa pudica causes
(a) t he plant t o bend t owar d light .
(b) st omat al opening so phot osynt hesis can begin.
(c) st r ess t hat r esult s in t he pr oduct ion of PR pr ot eins.
(d) leaves t o dr oop.
35. A biological cycle wit h a per iod of about 24 hour s is called
(a) t higmot r opism. (b) a cir cadian r hyt hm.
(c) phot oper iod. (d) abscission.
36. An Rajast hani hunt er was wor r ied about being at t acked by gr izzly t iger s, so he left
t he light s in his cabin on all t he t ime. Plant s near t he cabin flower ed a mont h ear ly.
Which of t he following best explains t his?
(a) They must have been long-night plant s.
(b) The light s must have emit t ed far -r ed light .
(c) They must have been long-day plant s.
(d) They must have been shor t -day plant s.
37. Most plant s flower when
(a) t he soil r eaches a cer t ain t emper at ur e.
(b) t he days ar e of r ight lengt h.
(c) a cer t ain number of days have passed since t hey last flower ed.
(d) t he night s ar e of r ight lengt h.
38. A cer t ain shor t -day plant flower s when days ar e less t han 12 hour s long. Which of t he
following would cause it t o flower ?
(a) a 9-hour night and 15-hour day wit h 1 minut e of dar kness aft er 7 hour .
(b) an 8-hr day and 16-hour night wit h a flash of whit e light aft er 8 hr .
(c) a 13-hour night and 11-hour day wit h 1 minut e of dar kness aft er 6 hr .
(d) a 12-hour day and 12-hour night wit h a flash of r ed light aft er 6 hr .
Practice Test Paper–Physiology 167
39. A chemical change in a subst ance called phyt ochr ome
(a) causes a plant t o bend t owar d light .
(b) t r igger s fr uit dr op.
(c) enables a plant t o r espond t o t he pr esence of light .
(d) is r esponsible for gr avit r opism.
40. Which of t he following t issues pr oduces volunt ar y body movement s?
(a) smoot h muscle (b) simple cuboidal epit helium
(c) car diac muscle (d) skelet al muscle
41. Which of t he following best illust r at es homeost asis?
(a) All t he cells in t he body have much t he same chemical composit ion.
(b) Cells of t he skin ar e const ant ly wor n off and r eplaced.
(c) When blood CO
2
incr eases, you br eat he fast er and get r id of CO
2
.
(d) All or gans ar e composed of t he same four kinds of t issues.
42. How does a gast r ovascular cavit y differ fr om an aliment ar y canal? The gast r ovascular
cavit y
(a) st or es food but does not digest it .
(b) absor bs food molecules but does not pr oduce hydr olyt ic enzymes.
(c) has only a single opening.
(d) funct ions in digest ion but not absor pt ion.
43. In humans, most nut r ient molecules ar e absor bed by t he
(a) st omach. (b) liver .
(c) small int est ine. (d) lar ge int est ine.
44. The lar gest var iet y of digest ive enzymes funct ion in t he
(a) lar ge int est ine. (b) or al cavit y.
(c) st omach. (d) small int est ine.
45. Which of t he following is not an essent ial nut r ient for a human?
(a) linoleic acid, a fat t y acid (b) glucose, a monosacchar ide
(c) met hionine, an amino acid (d) ascor bic acid, a vit amin
46. Individuals lacking adequat e levels of ent er ogast r one would have t he gr eat est difficult y
digest ing
(a) fat s. (b) pr ot eins.
(c) car bohydr at e (d) nucleic acids.
47. It is impor t ant t o get some vit amin B-1 ever y day, but it is all r ight if int ake of
vit amin A var ies a bit . Why?
(a) Vit amin B-1 is an essent ial nut r ient , and vit amin A is not .
(b) Vit amin A can be st or ed by t he body, but vit amin B-1 cannot .
(c) The body needs much lar ger amount s of vit amin B-1 t han vit amin A.
(d) The body r equir es vit amin B-1, but vit amin A is just an “ext r a.”
168 CSIR-NET Life Sciences
48. How would you expect t he digest ive syst em of a hawk, a car nivor e, t o compar e wit h
t hat of a spar r ow, a seed-eat er ?
(a) The hawk would have a lar ger gast r ovascular cavit y.
(b) The spar r ow’s digest ive syst em would be longer .
(c) The hawk would have a gizzar d, but t he spar r ow would not .
(d) The hawk digest ive syst em would be longer .
49. Gallst one sur ger y somet imes r equir es t hat t he gallbladder be r emove. Pat ient s ar e
t hen advised t o avoid ingest ing lar ge amount of fat because
(a) t he gallbladder makes bile, which is necessar y for fat emulsificat ion.
(b) wit hout t he bile pr oduced by t he gallbladder , fat s cannot be enzymat ically
hydr olyzed.
(c) t he gallbladder pr oduces t he hor mone ent er ogast r one.
(d) t he gallbladder st or es lar ge quant it ies of bile, r eleasing it when necessar y.
50. Resear cher s pr ovided r adioact ively labeled food t o a dog, and t r aced t he movement of
absor bed molecules. Which t ype of molecule would move along a differ ent pat h t han
all t he ot her s?
(a) car bohydr at es (b) pr ot eins
(c) nucleic acids (d) fat s
51. The car diac sphinct er sur r ounds t he car diac or ifice. If t his sphinct er failed t o pr oper ly
const r ict , t her e might be a pr oblem wit h
(a) r egur git at ion of food int o t he esophagus.
(b) movement of t he bolus int o t he t r achea r at her t han t he esophagus.
(c) r apid empt ying fr om t he st omach t o t he small int est ine.
(d) r apid empt ying fr om t he small int est ine t o t he lar ge int est ine.
52. Which of t he following might make t he most effect ive ant iulcer medicat ion? A chemical
t hat
(a) st imulat es par iet al cells of t he gast r ic glands.
(b) kills bact er ia in t he st omach.
(c) inhibit s mucous cells of t he gast r ic glands.
(d) st imulat es chief cells of t he gast r ic pit s.
53. The lungs consist of many small air sacs and blood vessels, which gr eat ly incr ease
sur fa ce a r ea a nd impr ove t he t r a nsfer of subst a nces t hr ough t heir wa lls. The
st r uct ur es in t he digest ive syst em similar in funct ion t o t hese air sacs and capillar ies
ar e t he
(a) villi. (b) colon and r ect um.
(c) gast r ic glands. (d) high-densit y lipopr ot eins.
54. Dur ing some t ypes of ant ibiot ic t r eat ment s, pat ient s oft en exper ience diar r hoea
because
(a) ant ibiot ics ar e t oxic t o t he colon’s epit helium as well as t o bact er ia.
(b) t he bact er ial flor a of t he lar ge int est ine digest fiber , which ot her wise would
cr eat e osmot ic pr essur e and r esult in decr eased wat er r eabsor pt ion.
Practice Test Paper–Physiology 169
(c) ant ibiot ics int er fer e wit h t he vit amin absor pt ion pr ocess nor mally occur r ing
wit hin t he lar ge int est ine.
(d) aft er int est inal bact er ia have been killed, an unusually lar ge amount of wat er is
r eabsor bed.
55. Imagine t hat you have eat en a meal cont aining t he following nut r ient s. Which would
not have t o be digest ed befor e being absor bed?
(a) pr ot ein (b) polysacchar ide
(c) disacchar ide (d) amino acid
56. Which of t he following best descr ibes an ar t er y?
(a) car r ies blood away fr om t he hear t
(b) car r ies oxygenat ed blood
(c) cont ains valves
(d) has t hin walls
57. In a fish, blood cir culat es t hr ough ____, while in a mammal, it cir culat es t hr ough
____.
(a) t wo cir cuit s . . . four cir cuit s (b) one cir cuit . . . t wo cir cuit s
(c) four cir cuit s . . . t wo cir cuit s (d) one cir cuit . . . four cir cuit s
58. A r ecor ding of t he elect r ical act ivit y of a pat ient ’s hear t shows t hat t he at r ia ar e
cont r act ing r egular ly and nor mally, but ever y few beat s t he vent r icles fail t o cont r act .
Which of t he following is pr obably funct ioning impr oper ly?
(a) AV node (b) semilunar valve
(c) cor onar y ar t er y (d) pacemaker
59. St r oke occur s when
(a) t he pacemaker becomes defect ive, pr oducing an ir r egular hear t beat .
(b) a blood clot ent er s and blocks one of t he cor onar y ar t er ies.
(c) a blood clot dislodges fr om a vein and moves int o t he lung, wher e it blocks a
pulmonar y ar t er y.
(d) a blood clot ent er s t he cer ebr al cir culat ion, blocking an ar t er y and causing t he
deat h of br ain t issue.
60. An advant age of gas exchange in wat er , compar ed wit h gas exchange in air , is t hat
(a) wat er usually cont ains a higher concent r at ion of oxygen t han air .
(b) wat er is easier t o move over t he r espir at or y sur face.
(c) t he r espir at or y sur face does not dr y out in wat er .
(d) vent ilat ion r equir es less ener gy in wat er .
61. In t he blood, bicar bonat e ions
(a) help t r anspor t oxygen.
(b) act as buffer s t o guar d against pH changes.
(c) ar e t r anspor t ed by hemoglobin.
(d) at t ach t o numer ous car bon dioxide molecules, keeping t hem fr om solut ion.
170 CSIR-NET Life Sciences
62. Most oxygen is car r ied by t he blood __. Most car bon dioxide is car r ied by t he blood__.
(a) at t ached t o hemoglobin . . . in t he for m of bicar bonat e ions
(b) dissolved in t he plasma . . . dissolved in t he plasma
(c) in t he for m of H+ ions . . . in t he for m of bicar bonat e ions
(d) at t ached t o hemoglobin . . . at t ached t o hemoglobin
63. Which of t he following is an endot her m?
(a) mouse (b) iguana
(c) fr og (d) t r out
64. As filt r at e passes t hr ough t he long loop of Henle, salt is r emoved and concent r at ed in
t he int er st it ial fluid of t he kidney medulla. (a)Because of t his high salt concent r at ion,
t he nephr on is able t o
(a) excr et e t he maximum amount of salt .
(b) neut r alize t oxins t hat might accumulat e in t he kidney.
(c) cont r ol t he pH of t he int er st it ial fluid.
(d) est ablish a hyper t onic int er st it ial medullar y concent r at ion.
65. Ur ic acid is t he nit r ogenous wast e excr et ed by bir ds, insect s, and many r ept iles. An
advant age of excr et ing ur ic acid is t hat it ____ , but a disadvant age is t hat it _____ .
(a) saves wat er . . . cost s ener gy
(b) saves ener gy . . . is highly t oxic
(c) is not ver y t oxic . . . wast es a lot of wat er
(d) is much mor e soluble in wat er t han ot her wast es . . . cost s ener gy
66. On a cold day, blood vessels in t he skin
(a) dilat e, allowing blood t o keep t he skin war m.
(b) const r ict , for cing blood t o flow t hr ough vessels in t he skin.
(c) const r ict , r educing heat loss fr om blood at t he sur face.
(d) dilat e, causing blood t o pass t hr ough t he cold skin mor e quickly.
67. The animals in which of t hese pair s have similar pr oblems r egulat ing wat er balance?
(a) fr eshwat er fish and salt wat er fish
(b) land animal and fr eshwat er fish
(c) osmoconfor mer and fr eshwat er fish
(d) salt wat er fish and land animal
68. Which would have t he t oughest t ime sur viving over t he long t er m in t he envir onment
given?
(a) an osmoconfor mer in seawat er
(b) an endot her m in a war m envir onment
(c) an ect ot her m in a cold envir onment
(d) an ect ot her m in a war m envir onment
Practice Test Paper–Physiology 171
69. Most aquat ic animals excr et e ammonia, while land animals excr et e ur ea or ur ic acid.
What is t he most likely explanat ion for t his differ ence?
(a) They have differ ent diet s.
(b) Land animals can get t he ener gy needed t o make ur ea or ur ic acid.
(c) Ammonia is ver y t oxic, and it t akes lot s of wat er t o dilut e it .
(d) Land animals cannot affor d t he ener gy needed t o make ammonia.
70. Which of t he following hor mones has t he br oadest r ange of t ar get s?
(a) ADH (b) TSH
(c) epinephr ine (d) ACTH
71. Ever y t ime you eat a cookie or candy bar , your blood sugar incr eases. This t r igger s
an incr ease in t he hor mone
(a) insulin (b) epinephr ine.
(c) adr enocor t icot r opin(ACTH). (d) glucagon.
72. Ever y hor mone
(a) is a pr ot ein.
(b) is pr oduced in r esponse t o st r ess.
(c) is under t he cont r ol of t he pit uit ar y gland.
(d) ent er s a cell and int er act s wit h DNA.
73. Since most chemical signals ar e unable t o pass t hr ough t he plasma membr ane, t he
cellular act ion t hey init iat e r esult s fr om
(a) ligand binding.
(b) t he act ivat ion of a signal t r ansduct ion pat hway.
(c) dir ect st imulat ion of t he cell’s DNA.
(d) t he enzymat ic behaviour of t he signal molecule.
74. Which of t he following hor mones have ant agonist ic (opposing) effect s?
(a) t hyr oxin and calcit onin
(b) insulin and glucagon
(c) gr owt h hor mone and epinephr ine
(d) ACTH and glucocor t icoids
75. What is t he r ole of a second messenger in hor mone act ion?
(a) It signals a cell t o secr et e a hor mone.
(b) It infor ms a gland as t o whet her it s hor mones ar e having an effect .
(c) It r elays a hor mone’s message inside a t ar get cell.
(d) It st ops hor mone act ion when it is no longer needed.
76. When t he levels of juvenile hor mone (J H) ar e maint ained at ar t ificially high levels,
insect s will
(a) be unable t o molt .
(b) bypass some lar val st ages and pupat e pr emat ur ely.
(c) molt mor e fr equent ly.
(d) be unable t o advance t o a pupal st age.
172 CSIR-NET Life Sciences
77. The act ion of a local r egulat or is illust r at ed when
(a) pr ost aglandins r eleased fr om t he placent a alt er t he excit abilit y of t he muscle of
t he ut er us.
(b) hist amine pr omot es t he secr et ion of hydr ochlor ic acid by st omach cells.
(c) t he r elaxat ion of ar t er ial smoot h muscle caused by nit r ic oxide.
(d) all of t he above.
78. Some glands pr oduce hor mones t hat st imulat e ot her endocr ine glands. Which of t he
following hor mones specifically act s t o t r igger secr et ion of hor mones by anot her
endocr ine gland?
(a) t hyr oid hor mone (T3 and T4)
(b) pr ogest er one
(c) adr enocor t icot r opin (ACTH)
(d) ant idiur et ic hor mone (ADH)
79. How is t he level of t hyr oxin in t he blood r egulat ed?
(a) Thyr oxin st imulat es t he pit uit ar y t o secr et e t hyr oid-st imulat ing hor mone (TSH).
(b) TSH inhibit s secr et ion of t hyr oxin fr om t he t hyr oid gland.
(c) Thyr oxin st imulat es t he hypot halamus t o secr et e TRH.
(d) Thyr oxin and TSH inhibit secr et ion of TRH.
80. The endocr ine and ner vous syst ems shar e which of t he following in common?
(a) Bot h ut ilize feedback.
(b) Sever al chemicals ser ve as bot h hor mones and neur ot r ansmit t er s.
(c) Ner ve impulses can cause endocr ine glands t o r elease hor mone.
(d) all of t he above.
81. It t akes much longer for sex hor mones and ot her st er oids t o pr oduce t heir effect s
t han it t akes nonst er oid hor mones. Why?
(a) St er oids ar e bigger , slower molecules.
(b) St er oids usually must be car r ied longer dist ances by t he blood.
(c) St er oids cause t ar get cells t o make new pr ot eins, which t akes t ime.
(d) St er oids must r elay t heir message via a second messenger .
82. Which disor der is cor r ect ly mat ched wit h it s cause?
(a) pit uit ar y dwar fism and hyposecr et ion of gr owt h hor mone
(b) infant cr et inism and hyper secr et ion of t hyr oxin
(c) low blood calcium and hyper secr et ion of par at hyr oid hor mone (PTH)
(d) acr omegaly and hyposecr et ion of gr owt h hor mone
83. Inject ions of a hor mone ar e somet imes given t o st r engt hen cont r act ions of t he ut er us
dur ing childbir t h. What hor mone might t his be?
(a) adr enocor t icot r opin (ACTH)
(b) t hyr oxin
(c) oxyt ocin
(d) follicle-st imulat ing hor mone (FSH)
Practice Test Paper–Physiology 173
84. Type I diabet es mellit us
(a) an aut oimmune disease t hat develops following immune at t ack on pancr eat ic
cells.
(b) a commonly seen in over weight individuals older t han for t y.
(c) is t r eat ed by impr oving insulin r ecept or efficiency r at her t han by giving insulin.
(d) is t he most common for m of t he disease; mor e t han 90% of all diabet ics have
t ype I.
85. J et lag occur s when a per son moves r apidly fr om one t ime zone t o anot her , causing
conflict bet ween t he body’s biological r hyt hm and t he new cycle of light and dar k.
Some scient ist s suspect t hat jet lag may r esult fr om disr upt ion of a daily hor mone
cycle. Which of t he following hor mones do you t hink is t he most likely suspect ?
(a) epinephr ine (b) insulin
(c) melat onin (d) est r ogen
86. As a young gir l, Sunit a suffer ed a head injur y t hat damaged her pit uit ar y. An injur y
t o t he pit uit ar y is par t icular ly ser ious because of all t he funct ions cont r olled by t his
gland As Sunit a got older , she and her doct or s found t hat all of t he following except
_____ wer e affect ed.
(a) met abolic r at e (b) blood sugar level
(c) her menst r ual cycle (d) milk pr oduct ion
87. A hor mone fr om t he par at hyr oid gland wor ks in opposit ion t o a hor mone fr om t he
_____ t o r egula t e _____ .
(a) post er ior pit uit ar y . . . met abolic r at e
(b) t hyr oid gland . . . blood calcium
(c) pancr eas . . . wat er r eabsor pt ion
(d) adr enal medulla . . . blood calcium
88. Diabet es insipidus is an inher it ed endocr ine malfunct ion (unr elat ed t o diabet es
mellit us) in which t he kidneys fail t o r eabsor b nor mal amount s of wat er . Vict ims of
t his disease pr oduce gallons of ur ine each day, and t heir kidneys soon wear out .
Tr eat ment of t his disease involves r eplacing a missing hor mone. Which of t he following
do you t hink it is?
(a) glucagon (b) epinephr ine
(c) glucocor t icoids (d) ant idiur et ic hor mone (ADH)
89. Because only t he _____________ gland uses iodine t o make it s hor mones, r adioact ive
iodine is oft en used as a t r eat ment for t umor s of t his gland.
(a) pit uit ar y (b) pancr eat ic
(c) t hyr oid (d) adr enal
90. Which of t he following is a for m of sexual r epr oduct ion?
(a) budding (b) fission
(c) fr agment at ion (d) her maphr odit ism
174 CSIR-NET Life Sciences
91. A peak in ______ t r igger s ovulat ion on about t he ______ day of t he mont hly cycle.
(a) LH . . . sevent h
(b) FSH . . . second
(c) LH . . . four t eent h
(d) est r ogen . . . t went iet h
92. Ext er nal fer t ilizat ion occur s most ly in
(a) land animals.
(b) insect s.
(c) aquat ic animals.
(d) animals t hat r epr oduce asexually.
93. On it s way t o fer t ilize a human egg, a sper m cell does not have t o pass t hr ough which
of t he following?
(a) oviduct (b) vagina
(c) ovar y (d) vas defer ens
94. Which of t he following hor mones is t he fir st t o incr ease significant ly ever y 28 days or
so and init iat es t he ovar ian cycle?
(a) pr ogest er one
(b) follicle-st imulat ing hor mone (FSH)
(c) est r ogen
(d) lut enizing hor mone (LH)
95. Aft er ovulat ion occur s, t he empt y follicle
(a) can be r ecycled t o pr oduce mor e eggs.
(b) changes int o t he cor pus lut eum and makes hor mones.
(c) quickly degener at es.
(d) immediat ely init iat es menst r uat ion.
96. Oogenesis in compar ison t o sper mat ogenesis is somewhat unusual in humans in t hat
(a) cyt okinesis is unequal dur ing t he meiot ic divisions.
(b) t he sequence fr om pr imar y oocyt e t o ovum is int er r upt ed by a r elat ively long
r est ing per iod.
(c) t he fir st meiot ic division is not complet ed unless r eact ivat ed by a hor mone.
(d) all of t he above
97. Bir t h cont r ol pills cont ain synt het ic est r ogen and pr ogest er one. How might t hese
hor mones pr event pr egnancy?
(a) They t r igger pr emat ur e ovulat ion, befor e an egg is mat ur e.
(b) They cause t he lining of t he ut er us t o be sloughed off.
(c) They cause t he cor pus lut eum t o degener at e.
(d) They keep t he pit uit ar y fr om secr et ing FSH and LH, so ovulat ion does not
occur .
Practice Test Paper–Physiology 175
98. Pr egnancy t est s det ect a hor mone hCG in a woman’s ur ine t hat is pr esent only when
an embr yo is developing in her ut er us. This hor mone is secr et ed by
(a) t he ovar y. (b) t he embr yo.
(c) a follicle. (d) t he endomet r ium.
99. In many mammals t he t est es ar e locat ed out side t he abdominal cavit y wit hin t he
scr ot um because
(a) t he elevat ed pr essur e wit hin t he abdominal cavit y would collapse t he small
passage ways wit hin t he t est es.
(b) t his locat ion allows for a shor t er pat hway t o t he ur et hr a.
(c) blood flow t o t he scr ot um is not int er r upt ed dur ing er ect ion.
(d) sper m ar e unable t o pr oper ly mat ur e at t he higher t emper at ur es found wit hin
t he abdominal cavit y.
100. Because t he genet ic composit ion of t he foet us is not ident ical t o t hat of t he mot her , it
is somewhat sur pr ising t hat t he foet us is not r eject ed as a for eign body. It appear s
t hat t his is because
(a) t he embr yo does not expr esses pat er nal ant igens on it s cells unt il aft er bir t h.
(b) t he embr yo pr oduces signal molecules t hat t ur n off t he mot her ’s immune syst em
for t he 9 mont hs of pr egnancy.
(c) a pr ot ect ive layer , t he t r ophoblast , sur r ounds t he embr yo and pr event s dir ect
cont act wit h mat er nal t issue.
(d) special mat er nal immune cells pr oduce ant igenic modifier pr ot eins (AMPs), which
mask t he for eign ant igens.
Practice Test Paper-II
1. One ear ly, and inexpensive, appr oach t o infer t ilit y is t o t est a woman’s ur ine for a
hor mone t hat would indicat e a high pr obabilit y of ovulat ion. What hor mone is t his
t est kit designed t o det ect ?
(a) oxyt ocin
(b) follicle-st imulat ing hor mone (FSH)
(c) t est ost er one
(d) lut enizing hor mone (LH)
2. A sper m has sever al component s; t he funct ion of t he acr osome is
(a) pr oduce base t o neut r alize t he acidic envir onment of t he female r epr oduct ive
syst em.
(b) met abolize t he sugar s pr ovided for ener gy by t he semen.
(c) r elease an enzyme t hat br eaks down t he membr ane of t he ovum.
(d) pr opel t he sper m as t hey swim t hr ough t he fluid of t he female r epr oduct ive
t r act .
3. The dr ug RU486 was developed in Fr ance and is widely used in Eur ope as a met hod
of bir t h cont r ol. It s int r oduct ion t o t he Unit ed St at es has been cont r over sial, however ,
t he dr ug is t aken aft er sexual int er cour se. It blocks t he implant at ion of t he developing
blast ocyst . Opponent s of t he dr ug say it is an abor t ion pill. It s defender s point out
t hat it wor ks t he same way as
(a) t he familiar bir t h cont r ol pill used for t he last 30 year s.
(b) sper micidal foam.
(c) an int r aut er ine device (IUD).
(d) t ubal ligat ion.
4. Acet ylcholinest er ase is t he enzyme t hat degr ades acet ylcholine. What effect on ner ve
t r ansmission would occur following t he administ r at ion of a chemical t hat inhibit ed
acet ylcholinest er ase?
(a) no effect
(b) Synapt ic t r ansmission would be pr event ed; muscle par alysis would occur .
(c) It would be ident ical t o giving an anest het ic, but would last per manent ly.
(d) The pr e-synapt ic neur on would be inact ivat ed.
5. Anest het ics block pain by blocking t he t r ansmission of ner ve signals. Which of t hese
t hr ee chemicals might wor k as anest het ics?
(a) a chemical t hat pr event s t he opening of sodium channels in membr anes;
(b) a chemical t hat inhibit s t he enzymes t hat degr ade neur ot r ansmit t er s;
(c) a chemical t hat blocks neur ot r ansmit t er r ecept or s.
(d) a and c bot h
6. What is t he differ ence bet ween a neur on and a ner ve?
(a) One is sensor y in funct ion, t he ot her mot or .
(b) Ner ves ar e found only in t he cent r al ner vous syst em.
Practice Test Paper–Physiology 177
(c) They consist of differ ent number s of cells.
(d) Neur ons ar e made of whit e mat t er , ner ves of gr ay mat t er .
7. The fir st st age of embr yonic development is ______. This pr ocess pr oduces _______.
(a) gast r ulat ion . . . a t hr ee-layer ed embr yo
(b) gest at ion . . . a gast r ula
(c) ovulat ion . . . a zygot e
(d) cleavage . . . a hollow ball of cells
8. The funct ion of t he amnion in bir ds, r ept iles, and mammals is
(a) t o digest yolk and for m a net wor k of blood vessels t o dist r ibut e nut r ient s t o t he
embr yo.
(b) t o collect ur ic acid.
(c) t o cr eat e an aqueous envir onment in which development can occur .
(d) t o t r anspor t oxygen t o t he embr yo.
9. Up t o t he eight -cell st age, t he blast omer es of a mouse embr yo can each for m a complet e
embr yo if isolat ed. This indicat es t hat
(a) differ ent iat ion does not depend on cyt oplasmic det er minant s.
(b) t he mouse embr yo is st r ongly polar ized.
(c) only t he zygot e is t ot ipot ent .
(d) cyt oplasmic det er minant s ar e equally dist r ibut ed dur ing t he ear ly cleavage
divisions.
10. One differ ence bet ween t he blast ula and gast r ula st ages of development is t hat
(a) blast ula cells ar e mor e differ ent iat ed t han gast r ula cells.
(b) t her e ar e many mor e cells in a blast ula.
(c) t he blast ula consist s of mor e cell layer s.
(d) t her e is an opening fr om t he cavit y inside t he gast r ula t o t he out side.
11. Among t he following which disease is not r elat ed t o vit amin deficiency:
(a) Ost eomalacia (b) Xer opht halmia
(c) Mar asmus (d) Pellagr a
12. A pr ot ein deficiency disor der char act er ized by mat ch st ick legs, pr ot r uded belly, oedema
of cer t ain par t s is:
(a) Mar asmus (b) Kwashior kor
(c) Ricket s (d) Ost eomalacia
13. Vit amin which cause Per nicious anaemia, glossit is, loss of per ipher al sensat ion due
t o it s deficiency and also called as Er yt hr ocyt e Mat ur ing Fact or is-
(a) Vit B2 (b) Vit B6
(c) Cobalt amine (d) Folic Acid
14. The t ype of anemia caused by deficiency of folic acid is-
(a) Megablast ic (b) Macr ocyt ic
(c) Per nicious (d) Thalassemia
178 CSIR-NET Life Sciences
15. Among t he following t he fat soluble vit amin is
(a) Folic Acid (b) Calcifer ol
(c) Ascor bic Acid (d) Biot in
16. Vit amin which is ant i-st er ilit y fact or , ant i-oxidat ive for membr ane lipids, skin and
hair r educes at her oscler osis is:
(a) Tocopher ol (b) Calcifer ol
(d) Phylloquinone (d) Pyr idoxine
17. Among t he following essent ial aminoacids is:
(a) Ar ginine (b) Hist idine
(c) Tyr osine (d) Glut amine
18. Which miner al is r equir ed for blood clot t ing, muscle cont r act ion, ATPase, ner ve
impulse t r ansmission, hear t funct ioning et c:
(a) Mg (b) Ca
(c) K (d) Fe
19. Miner al engaged in maint aining enamel and checking dent al decay is:
(a) Selenium (b) Pot assium
(c) Calcium (d) Fluor ine
20. Among t he following, which ar e not pept ide hor mone:
(a) ARH, MSH (b) Oxyt ocin, ACTH
(c) Calcit onin, , ADH (d) TSH, LTH, FSH
21. Relaxin is secr et ed by:
(a) Ant er ior Pit uit ar y (b) Post er ior pit uit ar y
(c) Cor pus Lut eum (d) Gr afian follicle
22. Which among t he following is not a funct ion of par at hor mone for maint aining opt imum
level of Ca
(a) By mobilizat ion fr om bone
(b) Reduced ur inar y excr et ion
(c) Incr eased absor pt ion fr om int est ine
(d) By mobilizat ion fr om muscles
23. Which among t he following is gast r o-int est inal hor mone:
(a) Insulin (b) Glucagons
(c) Ser ot onin (d) Secr et in
24. Dor mit or y effect (synchr onizat ion of menst r ual cycle of women living t oget her ) in
females is believed t o be due t o-
(a) Est r ogen (b) Pr ogest er one
(c) Pher omone (d) Oxyt ocin
25. Female cont r acept ive pills such as Mala-D gener ally cont ains :
(a) Pr ogest er one (b) Est r ogen
(c) Bot h a & b (d) hCG
Practice Test Paper–Physiology 179
26. Complet e (Obligat e) Par t henogenesis (males complet ely absent ) is pr esent in:
(a) Lacer t a saxicola ar meniaca (Caucasian r ock lizar d)
(b) Typhilina br ahmina (Small lizar d)
(c) Cnemidophor us (Whipt ail lizar d of Amer ica)
(d) All t he above
27. The cells which funct ions as nur se cells for differ ent iat ing sper mat ozoa:
(a) Leydig cells (b) Int er st it ial cells
(c) Ser t oli cells (d) Copus Epididymus
28. The t ype of egg/ovum of higher mammals on basis of amount of yolk is-
(a) Alecit hal (b) Micr olecit hal
(c) Macr olecit hal (d) Mesolecit hal
29. Glands of male r epr oduct ive syst em ar e:
(a) Pr ost r at e, seminal vesicle , mammar y
(b) Pr ost r at e, Ber t holin’s, seminal vesicle
(c) Seminal vesicle, Ber t holin’s gland
(d) Pr ost r at e, seminal vesicle, Cowper ’s
30. Temper at ur e in Scor t um necessar y for sper m for mat ion should be:
(a) 2 °C Above body t emper at ur e
(b) 2 °C Below body t emper at ur e
(c) 4 °C Above body t emper at ur e
(d) 4 °C Below body t emper at ur e
31. The Pr ocess of differ ent iat ion of a sper mat ozoa fr om a sper mat id is known as:
(a) Sper miogenesis (b) Sper mat ogenesis
(c) Sper mat ocyt ogeneis (d) Sper mat oneogenesis
32. Dur ing fer t ilizat ion compat abilt y r eact ion t akes place bet ween:
(a) Fer t ilizin (fr om ovum)& ant ifer t ilizin (Sper m)
(b) Fer t ilizin (fr om sper m)& ant ifer t ilizin (ovum)
(c) Fer t ilizin (sper m head)& ant ifer t ilizin (Sper m t ail)
(d) Fer t ilizin (sper m t ail)& ant ifer t ilizin (Sper m head)
33. Zygot e for mat ion by sper m and egg is r esult of phenomenon called-
(a) Fusion (b) Endocyt osis
(c) Phagocyt osis (d) En-nucleat ion
34. Which hor mone is called as st r ess hor mone:
(a) Auxin (b) Epinephr ine
(c) Et hylene (d) Nit r ous oxide
35. Which disease is not r elat ed t o t hyr oid gland:
(a) Gr ave’s disease (b) Hashimot o disease
(c) Addison disease (d) Cr et inism
180 CSIR-NET Life Sciences
36. Among t he following which is not cor r ect ly mat ched-
(a) cAMP-Pr ot ein Kinase A Pat hway
(b) IP
3
/DAG-Pr ot ein Kinase C Pat hway
(c) cGMP-Pr ot ein Kinase G Pat hway
(d) None of t hese
37. Among t he following which pair of hor mone is secr et ed by post er ior pit uit ar y
(a) FSH & LH (b) ADH & Oxyt ocin
(c) GH and PRL (d) ACTH and TSH
38. Among t he following shor t est pept ide hor mone is-
(a) Thyr ot r opin r eleasing hor mone
(b) Gonadot r opin r eleasing hor mone
(c) ACTH
(d) Insulin
39. Among t he following, which hor mone act s on ser t oli cells of seminifer ous t ubules t o
incr ease pr ot ein in sper m and ot her pr ot eins; act s on ovar ian follicle t o st imulat e t he
mat ur at ion of ovum and pr oduct ion of est r adiol-
(a) FSH (b) LH
(c) hCG (d) Pr ogest r one
40. Which hor mone st imulat es pancr eat ic acinar cells t o r elease bicar bonat e and wat er
in duodenum at pH values below 4.5 t o elevat e duodenal pH
(a) cholecyst okinin (CCK) (b) Gast r in
(c) Secr et in (d) Er yt hr opoeit in
41. Among t he following which is not secr et ed by ovar ian cor pus lut eum-
(a) Relaxin (b) est r ogen
(c) Human placent al lact ogen (hPL) (d) Pr ogest r one
42. Which hor mone st imulat es bone r esor pt ion; st imulat es phosphat e excr et ion by kidney
and r aises ser um calcium level-
(a) Calcit onin (b) Ser ot onin
(c) Pa r a t hor mone (d) Endot helin
43. Epinephr ine is synt hesized fr om amino acids-
(a) Phenyl allanine/Tyr osine (b) Met hionine/cyst eine
(c) Leucine/Isoleucine (d) Glycine/Pr oline
44. Wit h ant er ior pit uit ar y hor mones TSH, LH and FSH-
(a) a-subunit s ar e all differ ent
(b) b-subunit s ar e specifically r ecognized by r ecept or s
(c) b-unit alone can bind t o r ecept or
(d) int r acellular r ecept or s r ecognize t hese hor mones
Practice Test Paper–Physiology 181
45. St er oid hor mones ar e synt hesized fr om-
(a) Fat t y acids (b) St er ols
(c) Amino acids (d) Alcohols
46. The C-21 st er oid hor mone include-
(a) aldost er one (b) est r adiol
(c) t est ost er one (d) Vit amin D
3
47. Among t he following which hor mone binds t o t he membr ane r ecept or s-
(a) Thyr oxin (b) Est r ogen
(c) insulin (d) t est ost er one
48. Among t he following which pr ot ein is not involved in phot o t r ansduct ion cascade
dur ing r ecept ion of light by r od cells
(a) Rhodopsin (b) Tr anducin
(c) Phosphodiest r ase (d) Adenylat e cyclase
49. Among t he following which is not an t ype of excit at or y neur ot r ansmit t er in ner vous
t issue-
(a) Acet yl choline (b) Dopamine
(c) Hist amine (d) g-aminobut yr ic acid (GABA)
50. Thick filament of muscle is-
(a) Act in (b) Myosin
(c) Tr opomyosin (d) Tr oponin
51. Classic hemophelia is due t o deficiency of –
(a) Fact or IV (b) Fact or VIII
(c) Fact or IX (d) Fact or XI
52. The Effect of Vit amin A may include all of t he following except -
(a) Pr event ion of anemia (b) Ser ving as ant ioxidant
(c) The visual cycle (d) Induct ion of cer t ain cancer s
53. In t he pr opagat ion of ner ve impulse by an elect r ical signal-
(a) t he elect r ical pot ent ial acr oss t he membr ane is maint ained by ATP dr iven Na
+
-
K
+
pump becomes mor e negat ive
(b) local depolar izat ion of membr ane causes pr ot ein confor mat ional changes in ion
channels t hat allow Na
+
and K
+
t o move down concent r at ion gr adient
(c) Char ge pr opagat ion is bidir ect ional along axons
(d) “Volt age gat ed” ion channels have finit e r ecover y t ime so t he amplit ude of t he
impulse changes as it moves along t he axon.
54. ATP concent r at ion is maint ained r elat ively const ant dur ing muscle cont r act ion by-
(a) incr easing t he met abolic act ivit y
(b) t he act ion of adenylat e cyclase
(c) t he act ion of cr eat ine phosphokinase
(d) All t he above
182 CSIR-NET Life Sciences
55. An excessive int ake of calor ies-
(a) usually doesn’t have adver se met abolic consequences
(b) leads t o met abolic changes t hat ar e usually ir r ever sible
(c) fr equent ly leads t o elevat ed ser um level of fr ee fat t y acid, cholest er ol and t r i
acyl glycer ol
(d) is only component o obesit y
56. A complet e r eplacement of animal pr ot ein in diet by veget able pr ot ein
(a) would r educe t he t ot al amount of food consumed for t he same number of calor ies
(b) might r educe t ot al amount of Ir on and cobalt maine available
(c) Would be sat isfact or y r egar dless of nat ur e of veget able pr ot ein used
(d) Could not sat isfy pr ot ein r equir ement s
57. Among t he following, which is NOT a funct ion of t ocopher ol (Vit amin-E)
(a) Ant ioxidant (b) enhance heme synt hesis
(c) Pr event oxidat ion of LDL (d) car boxilat ion of pr ot hr ombin
58. Ascor bic acid (Vit amin C) may be associat ed wit h all of t he following EXCEPT-
(a) ir on absor pt ion
(b) bone for mat ion
(c) acut e r enal disease when t aken in high dose
(d) wound healing
59. St omat a opens when guar d cells-
(a) sense an incr ease in CO
2
in t he air spaces of leaf
(b) flap open because decr ease in t ur gor pr essur e
(c) become mor e t ur gid because of an influx of K
+
, followed by ent r y of wat er
(d) close aquapor ins, pr event ing wat er upt ake.
60. The movement of sap fr om a sugar sour ce t o sugar sink
(a) occur s t hr ough apoplast of sieve t ube member s
(b) may t r anslocat e sugar s fr om t he br eakdown of st or ed st ar ch in a r oot t ip t o
developing shoot s
(c) is similar t o flow of xylem sap in depending on t ension or negat ive pr essur e
(d) All of t he above
61. The pr oduct ivit y of cr op declines when leaves begin t o wilt mainly because-
(a) t he chlor oplast of wilt ing leaves decomposes
(b) flaccid mesophyll cells ar e incapable of phot osynt hesis
(c) st omat a close, pr event ing CO
2
fr om ent er ing t he leaf
(d) Phot olysis of wat er cannot occur
62. Micr onut r ient s ar e needed in ver y small amount because-
(a) most of t hem ar e mobile in plant s
(b) most funct ion as cofact or for enzymes
(c) most ar e supplied in lar ge quant it ies in seed
(d) only t he gr owing r egion of plant r equir e t hem
Practice Test Paper–Physiology 183
63. Mycor r hyzae enhance plant nut r it ion mainly by
(a) absor bing wat er and miner als t hr ough fungal hyphae
(b) conver t ing at mospher ic nit r ogen t o ammonia
(c) enabling t he r oot s t o par asit ize neighbor ing plant s st imulat ing t he development
of r oot hair s
(d) st imulat ing t he development of r oot hair s
64. Car nivor ous plant mainly compensat e for soil t hat has r elat ively low cont ent of-
(a) K
+
(b) Nit r ogen
(c) PO
4

(d) Ca
++
65. A miner al deficiency is likely t o affect older leaves mor e t hen younger leaves if
(a) t he miner al is micr onut r ient
(b) miner al is mobile wit hin plant
(c) miner al is r equir ed for chlor ophyll synt hesis
(d) Deficiency per sist s for longer t ime
66. The Thompson seedless gr ape is t r iploid, wit h t hr ee copies of each chr omosome.
Which phase of t he cell cycle would you expect t r iploid cells t o be unable t o complet e.
(a) meiosis 1 (b) S
(c) meiosis 2 (d) G2
67. Some or ganisms ar e capable of asexual or sexual r epr oduct ion. Under favor able
condit ions, r epr oduct ion pr oceeds asexually. When condit ions become mor e st r essful
r epr oduct ion swit ches t o a sexual mode. Why?
(a) Sexual r epr oduct ion is simple and mor e r apid allowing lar ger number s of offspr ing
t o be pr oduced.
(b) Sexual r epr oduct ion r equir es t wo separ at e individuals, who can mut ually pr ovide
nut r ient suppor t dur ing st r ess.
(c) Asexual r epr oduct ion r equir es mor e ener gy.
(d) Sexual r epr oduct ion pr oduces individuals wit h new combinat ions of r ecombined
chr omosomes incr easing diver sit y.
68. The st age of meiosis wher e cells become haploi(d)
(a) pr ophase I (b) pr ophase II
(c) anaphase I (d) anaphase II
69. One of t he ear liest event s t hat dist inguishes meiosis occur s in pr ophase I and involves:
(a) Condensa t ion of chr omosomes
(b) Loss of t he nuclear membr ane
(c) Movement of chr omosomes t owar ds t he met aphase plat e
(d) Pa ir ing of homologous chr omosomes
70. Cor al in t he ocean gr ows by budding, wher e t he new or ganism gr ows out of t he old
one by mit osis. This for m of r eplicat ion is an example of:
(a) meiosis t o pr oduce a zygot e (b) asexual r epr oduct ion
(c) sexual r epr oduct ion (d) gamet e for mat ion
184 CSIR-NET Life Sciences
71. ________ most closely r esembles event s of mit osis except t hat t he cells ar e ________.
(a) int er phase, diploid (b) meiosis II, diploid
(c) int er phase, haploid (d) meiosis II, haploid
72. Which of t he following is not alive, but r equir es life t o be able t o r epr oduce?
(a) Eubact er ia (b) Fungae
(c) Pr ot ozoa (d) Vir uses
73. In plant s IAA causes cell elongat ion due t o-
(a) Incr ease in pH of Apoplast (b) Incr ease in pH of cyt oplasm
(c) Decr ease in pH of Apoplast (d) Incr ease in pH of cyt oplasm
74. A hor mone t hat cont r ols closur e of st omat a in r esponse t o wat er st r ess is-
(a) Abscissic acid (b) Gibber ellins
(c) Pr oline (d) Et hylene
75. Which of t he following plays t he impor t ant r ole in t he mat ur at ion of sper mat ocyt e
out side t he seminal vesicle of t est is-
(a) Leydig Cells (b) Ser t oli Cells
(c) Sper ms (d) Sper mat ocyt es
76. Among t he following which phyt ohor mone incr eases t he femaleness t endency in plant s-
(a) GA (b) IBA
(c) Et hylene (d) Kinet in
77. Which hor mone is r elat ed wit h á-amylase act ivit y in ger minat ing seeds-
(a) GA (b) Kinet in
(c) Auxin (d) ABA
78. The amino acid which help in osmo r egulat ion dur ing st r ess condit ion is-
(a) Pr oline (b) Alanine
(c) b-glycine (d) Bot h a&c
79. The plant s ar e t er med as halophyt es if t hey ar e gr owing in soil having salt concent r at ion
above-
(a) 0.1 % (b) 0.2 %
(c) 0.4 % (d) 0.5 %
80. CAM mechanism is obser ved in plant s having-
(a) wat er st r ess (b) Salt st r ess
(c) No st r ess (d) Oxygen st r ess
81. Which among t he following is not an endocr ine gland?
(a) Pineal (b) Pit uit ar y
(c) Adr enals (d) Gonads
82. The excr et a of lizar ds is r ich in
(a) ur ea (b) ur ic acid
(c) guanidine (d) alant oin
Practice Test Paper–Physiology 185
83. Which one of t he following suppor t s glycogen synt hesis
(a) High cyclic adenosine monophosphat e (cAMP) levels
(b) Inact ive adenyl cyclase
(c) Act ive phosphor ylase-a
(d) Epinephr ine
84. Which of t he following vit amins, if t aken in excess is least likely t o cause pr oblems in
a healt hy per son?
(a) Vit amin A (b) Vit amin B
(c) Vit amin C (d) Vit amin D
85. When a muscle cont r act s, what is happening t o t he Ca
++
levels inside and out side t he
cell?
(a) High amount s of cyt osolic Ca
++
ar e r eleased t o t he ext r acellular space
(b) Ion channels open t o allow ext r acellular Ca
++
t o flow int o t he cell
(c) Ca
++
fr om t he nucleus is r eleased t o t he cyt oplasm and t his t r igger s cont r act ion
(d) Ca
++
ions at t ach st oma and t his causes muscle cont r act ion
86. Na
+
-K
+
pump is-
(a) Sympor t syst em (b) Ant ipor t syst em
(c) ABC t r anspor t er (d) Diffusion pump
87. Consider t he following st at ement s. Which is cor r ect ?
(a) Int r acellular concent r at ion of Na
+
is always high
(b) Int r acellular concent r at ion of K
+
ion is low
(c) Ext r a cellula r concent r a t ion of K
+
a n d Na
+
ion is isot onic t o int r a cellula r
concent r at ion
(d) Ext r a cellular concent r at ion of Na
+
and int r acellular concent r at ion of K
+
is always
high in nor mal cell.
88. The nut r it ional medium was supplied t o gr owing bact er ia. If whole nut r ient get
deplet ed in 20 cycles, t hen st age at which t he amount of nut r ient was half used-
(a) At end of 10 cycle (b) At beginning of 19 cycle
(c) At end of 19 cycle (d) At end of 11 cycle
89. Phyt ohor mone r esponsible for conver sion of st or ed pr ot eins int o glucose in ger minat ing
cer eals is-
(a) Cyt okinin (b) Auxin
(c) Gibber ellin (d) Abscissic Acid
90. Dir ect oxidat ion of per oxides in plant s is car r ied our by-
(a) Super oxide Dismut ase (b) Glut at hione Synt hase
(c) Cat alase (d) Per oxidase
91. A st r ong pept ide bond is pr esent in ar ginine and aspar t ic acid. This Bond will be weak
if ar ginine is being r eplaced by-
(a) Glut amic Acid (b) Lysine
(c) Hist idine (d) Pr oline
186 CSIR-NET Life Sciences
92. Mainly absor pt ion of glucose by int est ine is by-
(a) Na-K Pumps (b) Diffusion
(c) Na-glucose Sympor t er s (d) Unipor t er s
93. Pepsin which digest pr ot ein donot digest t he cells of int est ine because-
(a) Int est ine cells do not have pr ot eins
(b) Half life of pepsin is ver y low
(c) Pepsin act s only in acidic pH
(d) Pepsin do not digest int est ine pr ot eins
94. Dur ing ner ve impulse when acet yl choline binds r ecept or s, t her e is-
(a) In flow of Na & K ions
(b) In flow of Na and out flow of K ions
(c) In flow of K ions and out flow of Na ions
(d) In flow of K and H ions
95. Cellulose digest er digest cellulose of fer ment at ion. For e gut fer ment or s such as r abit
and elephant fer ment befor e int est ine while hind gut digest or like deer fer ment in
lar ge int est ine. It suggest t hat -
(a) Hind gut fer ment or ar e effect ive digest or of Cellulose
(b) For e gut fer ment or ar e effect ive digest or of Cellulose
(c) Bot h hind gut & for e gut fer ment or ar e effect ive digest or of Cellulose
(d) Cellulose digest ion do not depend on gut posit ion
96. Addison disease and Cushing syndr ome ar e due t o malfunct ioning of –
(a) Adr enal medulla (b) Adr enal cor t ex
(c) Thyr oid (d) Pit uit ar y
97. Which hor mone is pr esent in high amount dur ing pr egnancy in ur ine and used for
pr egnancy t est -
(a) hCG (b) Pr ogest r one
(c) Est r ogen (d) Relaxin
98. Osmot ic pot ent ial in plant cell is maint ained by-
(a) Pr oline & bet a Glycine (b) Hist idine
(c) Lysine (d) Glycine
99. Sper ms ar e mor phologically fit but ar e unable t o swim due t o defect in-
(a) Sper mat ogenesis (b) Sper miogenesis
(c) Pr ost r at e glands (d) Epidydimis
100. Which hor mone play impor t ant r ole in wat er st r ess
(a) Absiccic acid (b) Et hylene
(c) Gibber lic acid (d) Auxin
4
GENETICS
THE FOUNDATION OF GENETICS
• Five t housand year s ago or ear lier , people wer e using applied genet ics in t he for m of
plant and animal br eeding. The foundat ion for t he science of genet ics was laid in
1866, when Gr egor Mendel used va r iet ies of pea s t o conduct exper iment s on
inher it ance. Mendel’s r esear ch was ignor ed unt il t he t ur n of t he t went iet h cent ur y.
Plant breeders showed that both parents contribute equally to inheritance
• Plant s have some desir able char act er ist ics for genet ic st udies.
• Plant s can be gr own in lar ge quant it ies. They pr oduce lar ge number s of offspr ing.
Many have bot h male and female r epr oduct ive or gans in t he same plant , which allows
self-fer t ilizat ion. The gener at ion t ime is r elat ively shor t . It is easy t o cont r ol which
individuals mat e.
• J osef Got t lieb Kölr eut er made a few obser vat ions t hat Mendel lat er found useful.
• Kölr eut er ’s r esear ch focused on t he r elat ive cont r ibut ion of males and females t o t he
genet ics of plant s. Recipr ocal cr osses helped pr ove t hat bot h male and female par ent s
cont r ibut e equally t o t he char act er ist ics inher it ed by offspr ing. An example is a cr oss
of a male whit e plant wit h a female pink plant , and t he r ecipr ocal cr oss of a male pink
wit h a female whit e plant . The r esult ing pr ogeny have t he same appear ance.
• Befor e t he accept ance of Mendel’s r esear ch, t he concept of blending was favor ed. An
example of t his for m of inher it ance would be pur ple pr ogeny r esult ing fr om r ed and
blue par ent s. As a consequence of blending, t he pur ple color would be for ever blended
and could not be separ at ed.
Mendel’s discoveries were overlooked for decades
• Gr egor Mendel was a monk wit h scient ific t r aining in mat hemat ics, physics, and
biology. Mendel pr esent ed his nine-year -long pr oject or ally in 1865 and in wr it ing in
188 CSIR-NET Life Sciences
1866. His dat a challenged t he blending concept , but he was ahead of his t ime. His
t heor y was ignor ed, per haps because his biological peer s wer e not accust omed t o
r eviewing mat hemat ical dat a. Even Dar win, whose evolut ionar y t heor y r est s on genet ic
var iat ion among individuals, failed t o under st and Mendel’s point and r elied on t he
blending concept .
• In 1900, Hugo de Vr ies, Kar l Cor r ens, and Er ich von Tscher mak each independent ly
published paper s on t he quant it at ive out comes fr om cr osses. Each cit ed Mendel’s
t hen r ediscover ed 1866 paper . By 1900, meiosis had been descr ibed and Mendel’s
ideas wer e finally accept ed.
MENDEL’S EXPERIMENTS AND THE LAWS OF INHERITANCE
Mendel devised a careful research plan
• Mendel chose gar den peas as his subject s as t hey ar e easily gr own and t heir pollinat ion
is easily cont r olled. He cont r olled pollinat ion by manually moving pollen bet ween
plant s. He could also allow t he plant s t o self-pollinat e (using emasculat ion and bagging).
Mendel examined var iet ies of peas for her it able char act er s and t r ait s for his st udy. A
char act er is a feat ur e, such as flower color . A t r ait is a par t icular for m of a char act er ,
such as whit e flower s. “Her it able” means t he t r ait can be passed fr om par ent t o
pr ogeny.
• Mendel looked for char act er s t hat had well-defined alt er nat ive t r ait s and t hat wer e
t r ue br eeding. A t r ait is t r ue br eeding when it is t he only t r ait t hat occur s t hr ough
many gener at ions of br eeding individuals t hat shar e t hat t r ait . A t r ue-br eeding whit e-
flower ed plant would have only whit e-flower ed pr ogeny when cr ossed wit h ot her s in
it s st r ain.
• Tr ue-br eeding plant s, when used for cr ossing wit h ot her plant s t hat have an alt er nat ive
t r ait , ar e called t he par ent al gener at ion, designat ed P. The pr ogeny fr om t he cr oss of
t he P par ent s ar e called t he fir st filial gener at ion, designat ed F
1
. When F
1
individuals
ar e cr ossed t o each ot her or self-fer t ilized, t heir pr ogeny ar e designat ed F
2
.
• Mendel’s well-or ganized plan allowed him t o obser ve and r ecor d t he t r ait s of each
gener at ion in sufficient quant it y t o explain t hr ough analysis t he r elat ive pr opor t ions
of t he kinds of pr ogeny. His paper is r ecognized t oday as a model of clar it y.

Genetics 189
Mendel’s Experiment 1 examined a monohybrid cross
• Mendel cr ossed t r ue-br eeding plant s t hat differ ed for a given char act er . A monohybr id
cr oss involves one (mono) char act er and differ ent (hybr id) t r ait s. Pollen fr om t r ue-
br eeding pea plant s wit h wr inkled seeds (one t r ait ) was placed on st igmas of t r ue-
br eeding plant s wit h spher ical seeds (anot her t r ait ).
• The F
1
seeds wer e all spher ical; t he wr inkled t r ait failed t o appear at all. Because t he
spher ical t r ait complet ely masks t he wr inkled t r ait when t r ue-br eeding plant s ar e
cr ossed, t he spher ical t r ait is called dominant, and t he wr inkled t r ait is called recessive.
• Th e F
1
plant s wer e allowed t o self-pollinat e. This st ep was t he monohybr id cr oss.
This is also called an F
1
cross. S elf-pollination is sometimes called sel f i n g.
• The pr ogeny, called F
2
, wer e examined: 5,474 wer e spher ical and 1,850 wer e wr inkled
in r at io 3:1
• Mendel pr oposed t hat t he unit s r esponsible for inher it ance wer e discr et e par t icles.
They exist ed wit hin an or ganism in pair s, separ at ed dur ing gamet e for mat ion, and
r et ained t heir int egr it y. This is called t he p a r t i cu l a t e t h e or y, which is in shar p
cont r ast t o t he blending t heor y. Each pea has t wo unit s of inher it ance for each
char act er . Dur ing pr oduct ion of gamet es, only one of t he pair member s for a given
char act er passes t o t he gamet e. When fer t ilizat ion occur s, t he zygot e get s one fr om
each par ent , r est or ing t he pair .
• Mendel’s unit s of inher it ance ar e now called gen es. Differ ent for ms of a gene ar e
called alleles. Each allele is given a symbol. In t he case of wr inkled seeds, S might
r epr esent smoot h and s wr inkled. By convent ion, upper case S r epr esent s t he dominant ;
lower ca se s r epr esent s t he r ecessive.
• Tr ue-br eeding individuals would have t wo copies of t he same allele. Wr inkled would
be ss (t wo copies of same allele = homozygous). Smoot h t r ue-br eeding would be S S
(t wo copies of same allele = homozygous). Some smoot h-seeded plant s could be S s,
alt hough t hey would not be t r ue-br eeding. Individuals t hat ar e smoot h, but had a
wr inkled par ent , ar e het erozygous: S s.
• When an or ganism is st udied for t hr ee differ ent genes and has t he alleles AABbCC, it
is homozygous for A and C genes but het er ozygous for t he B gene.
190 CSIR-NET Life Sciences
• The physical appear ance of an or ganism is it s phenotype. Wr inkled-seed would be a
phenot ype. The act ual composit ion of t he or ganism’s alleles for a gene is it s genotype:
S s is a genot ype.
• Organisms have many different genes—some have t housands, and complex organisms
have 10 t imes t hat number. Most of t hese genes are yet t o be described in t erms of t he
DNA sequence or t he amino acid sequence of t he gene product .
Mendel’s first law says that alleles segregate
• When an individual pr oduces gamet es, alleles separ at e, so each gamet e r eceives one
member of t he pair of alleles. Th i s i s Me n d e l ’s fi r s t l a w, t h e l a w of s e gr e ga t i on .
• When fer t ilizat ion occur s, pair s ar e r eest ablished by r eceiving one copy fr om each
par ent .
• The Punnet t squar e is a simple box-like device t hat helps us t o consider all genet ic
combinat ions and can pr ovide clar it y by showing t he expect ed fr equencies of genot ypes.
The S and s symbols r epr esent t he single allele each gamet e r eceives. Fer t ilizat ion
pr ovides t he t wo alleles for t he new individual, one fr om t he male (sper m) and one
fr om t he female (egg).
• The Punnet t squar e shows t hat t he genot ypes and associat ed r at ios for a monohybr id
cr oss ar e 1 S S : 2 S s :1 ss.
• Any pr ogeny wit h an S would have t he dominant (smoot h) phenot ype, so t he phenot ypic
r at io is 3 smoot h t o 1 wr inkled.
• Now it is known t hat a gene is a por t ion of t he chr omosomal DNA t hat r esides at a
par t icular sit e, called a locus (plur al is loci). The gene codes for a par t icular funct ion.
Mendel ar r ived at t he law of segr egat ion wit h no knowledge of meiosis or chr omosomes.
The mechanism of chr omosome separ at ion in me i os i s I t oday explains his law of
segr egat ion.
Genetics 191
Mendel verified his hypothesis by performing a test cross
• A t e s t cr os s can det er mine t he genot ype (heterozygous or homozygous) of an individual
wit h a dominant t r ait . It involves cr ossing t he individual t o a t r ue-br eeding r ecessive
(homozygous r ecessive).
• If t he unknown is het er ozygous, a ppr oxima t ely ha lf t he pr ogeny will ha ve t he
dominant t r ait and half t he r ecessive t r ait (Ra t i o 1:1).
• If t he unknown is homozygous dominant , all t he pr ogeny will have t he dominant
t r ait .
• Ba ck cr os s is cr oss of F1 wit h any one of t he par ent . Gener ally, back cr oss wit h
homozygous r ecessive pa r ent s is ma de in pla nt br eeding a pplica t ion t o ensur e
maximum homozygosit y in or der t o obt ain pur e line var iet y.
Deviation from Mendelian Monohybrid cross:
• While Mendel discussed t r ait s, we now know t hat genes ar e segment s of t he DNA
t hat code for specific pr ot eins. These pr ot eins ar e r esponsible for t he expr ession of
t he phenot ype. The basic pr inciples of segr egat ion and independent assor t ment as
wor ked out by Mendel ar e applicable even for sex-linked t r ait s.
• Cod omi n a n t a l l e l e s : Codominant alleles occur when r at her t han expr essing an
int er mediat e phenot ype, t he het er ozygot es expr ess bot h homozygous phenot ypes.
An example is in human ABO blood t ypes, t he het er ozygot e AB t ype manufact ur es
ant ibodies t o bot h A and B t ypes. Blood Type A people manufact ur e only ant i-B
ant ibodies, while t ype B people make only ant i-A ant ibodies. Codominant alleles ar e
bot h expr essed. Het er ozygot es for codominant alleles fully expr ess bot h alleles. Blood
t ype AB individuals pr oduce bot h A and B ant igens. Since neit her A nor B is dominant
over t he ot her and t hey ar e bot h dominant over O t hey ar e said t o be codominant .
• I n comp le t e d omi n a n ce : Incomplet e dominance is a condit ion when neit her allele
is domina nt over t he ot her . The condit ion is r ecognized by t he het er ozygot es
expr essing an int er mediat e phenot ype r elat ive t o t he par ent al phenot ypes. If a r ed
flower ed plant is cr ossed wit h a whit e flower ed one, t he pr ogeny will all be pink.
When pink is cr ossed wit h pink, t he pr ogeny ar e 1 r ed, 2 pink, and 1 whit e.
• Flower color in snapdr agons is an example of t his pat t er n. Cr oss a t r ue-br eeding r ed
st r ain wit h a t r ue-br eeding whit e st r ain and t he F1 ar e all pink (het er ozygot es). Self-
fer t ilize t he F1 and you get an F2 r at io of 1 r ed: 2 pink: 1 whit e. This would not
happen if t r ue blending had occur r ed (blending cannot explain t r ait s such as r ed or
whit e skipping a gener at ion and pink flower s cr ossed wit h pink flower s should pr oduce
ONLY pink flower s).
Mendel’s second law says that alleles of different genes assort independently
• The second law descr ibes t he out come of d i h yb r i d (t wo char act er ) cr osses, or hybr id
cr osses involving addit ional char act er s. A dihybr id is an individual t hat is a double
het er ozygot e (e.g., wit h t he genot ype S sYy).
• Mendel’s second law st at es t hat t he S s alleles assor t int o gamet es independent ly of
t he Yy alleles. The dihybr id, S sYy, pr oduces gamet es t hat have one allele of each
gene. Four differ ent gamet es ar e possible and will be pr oduced in equal pr opor t ions:
192 CSIR-NET Life Sciences
S Y, S y, sY, and sy. Random fer t ilizat ion of gamet es yields t he out come visible in t he
Punnet t squa r e. Not e it s 4× 4 t a ble const r uct ion t o a ccommoda t e 16 possible
phenot ypes. Filling in t he t able and adding t he like cells r eveals a 9:3:3:1 r at io of t he
four possible phenot ypes (smoot h yellow, smoot h gr een, wr inkled yellow, and wr inkled
gr een). Th e l a w of i n d e p e n d e n t a s s or t me n t st at es t hat alleles of differ ent genes
assor t independent ly of one anot her dur ing gamet e for mat ion.
• In fact , t his law is not always t r ue. The law of independent assor t ment is accur at e for
genes t hat ar e on separ at e chr omosomes, but not necessar ily for genes t hat ar e on
t he sa me chr omosome (d u e t o Li n k a ge).
• Genes t hat are close t o each ot her on t he same chromosome t end t o st ay t oget her, but
crossing over during meiosis may separat e t hem. The closer t oget her on t he same
chromosome genes are, t he more t hey t end t o st ay t oget her.
Punnett squares or probability calculations: A choice of methods
• Mu l t i p l yi n g p r ob a b i l i t i e s : If t wo coins, a penny and a dime, ar e t ossed, each act s
independent ly of t he ot her . The pr obabilit y of bot h landing on heads would be 1/2× 1/
2 = 1/4. To find t he pr obabilit y t hat independent event s will bot h happen, t he gener al
r ule is t o mult iply t he pr obabilit ies of t he individual event s.
• Mon oh yb r i d cr os s p r ob a b i l i t i e s : In t he example of smoot h and wr inkled seeds,
het er ozygot es pr oduce S and s gamet es. The pr obabilit y of a gamet e being S is 1/2.
The pr obabilit y of t hat an F
2
plant will be S S is 1/2× 1/2 = 1/4.
• Ad d i n g p r ob a b i l i t i e s : The pr obabilit y of an event t hat can occur in t wo or mor e
ways is t he sum of t he pr obabilit ies for each way in which t he event can occur . For
example, t he genot ype S s can r esult fr om s in t he egg and S in t he sper m, or fr om S
in t he egg and s in t he sper m. Thus, t he pr obabilit y of het er ozygot es in t he F
2
gener at ion of a monohybr id cr oss is 1/4 + 1/4 = 1/2.
• Th e d i h yb r i d cr oss: To calculat e t he pr obabilit ies of t he out comes of dihybr id cr osses,
simply mult iply t he out comes fr om each of t he individual monohybr id component s.
For example, t he pr obabilit y of t he S S Yy genot ype can be calculat ed as follows: An F
1
Genetics 193
(dihybr id) cr oss of S sYy gener at es 1/4 S S , 1/2 S s, 1/4ss, and 1/4 YY, 1/2 Yy, 1/4 yy. The
pr obabilit y of t he S S Yy genot ype is t he pr obabilit y of t he S S genot ype, which is 1/4,
t imes t he pr obabilit y of t he Yy genot ype, which is 1/2. This would be 1/8 (1/4× 1/2).
Mendel’s laws can be observed in human pedigrees
• Pat t er ns for over 2,500 inher it ed human char act er ist ics have so far been det er mined.
Humans cannot be st udied using planned cr osses, so human genet icist s r ely on
pedigr ees, which show phenot ype segr ega t ion in sever a l gener a t ions of r ela t ed
individuals. Since humans have such small number s of offspr ing, human pedigr ees
do not show clear pr opor t ions. The act ual number of affect ed ver sus unaffect ed
offspr ing is impossible t o pr edict for a cer t ain couple because out comes for small
samples fail t o follow closely t he expect ed out comes.
• If neit her par ent has a given phenot ype, but it shows up in pr ogeny, t he t r ait is
r ecessive and t he par ent s ar e het er ozygous. The chance of ot her childr en get t ing t he
t r ait is 1/4. Half of t he childr en fr om such a cr oss will be car r ier s (het er ozygous for
t he t r ait ). The pr obabilit y of a car r ier (het er ozygot e) for a r ar e allele unknowingly
mar r ying anot her unr elat ed car r ier is quit e low.
• F or a d omi n a n t a l l e l e, every affect ed per son has an affect ed par ent . About half of
t he offspr ing of an affect ed per son ar e also affect ed (assuming only one par ent is
affect ed). The phenot ype occur s equally in bot h sexes.
• F or a r a r e r e ce s s i ve t r a i t , Affect ed people usually have par ent s who ar e bot h not
affect ed. One-quar t er of t he childr en, on aver age, of unaffect ed par ent s would be
affect ed. The phenot ype occur s equally in bot h sexes.
• Mar r iage bet ween close r elat ives r esult s in a higher likelihood t hat bot h par ent s will
be car r ier s of a r ar e allele and pr oduce affect ed childr en. The major use of pedigr ee
analysis is in clinical evaluat ion and counseling of pat ient s wit h inher it ed abnor malit ies.
SOME EXCEPTIONS TO MENDEL’S LAWS
I. Linkage
If t wo differ ent genes ar e locat ed r elat ively close t o each ot her on t he same chr omosome t hey
can not segr egat e independent ly. Ter m ce n t i mor ga n (cM) is used in e u k a r yot i c ge n e t i cs
and ma p u n i t /Mor ga n in mi cr obi a l gen et i cs . The number s of linkage gr oups ar e equal t o
number of haploid set s of chr omosomes. But in XX-XY t ype of or ganisms, number of linkage
gr oups will be one mor e in male t han in t he female as in male X and Y bot h ar e differ ent
chr omosomes.
• Cis – tans a r r a n ge me n t of ge n e s : If t he dominant alleles A, B of t wo linked genes
ar e pr esent on t he same chr omosome and t heir r ecessive alleles ar e pr esent on t he
homologous chr omosomes, t he ar r angement of genes is called ci s -a r r a n ge me n t .
On t he ot her hand, if one dominant gene and ot her r ecessive gene ar e pr esent on t he
same chr omosome (A,b) and t heir alleles t ype (a,B) on t he homologous chr omosome,
t his t ype of ar r angement is called t r ans-ar r angement .
194 CSIR-NET Life Sciences
• Recombinat ion Fr equency = 100
pr ogeny Tot al
s r ecombinat of Number
×
• Sever al ascomycet es fungi (e.g., Neur ospor a) pr oduce asci, which hold t he haploid
ascospor es pr oduced aft er meiosis in a specific linear or der , an or der ed t et r ad. This
or der r eflect s t he or ganizat ion and chr omat ids involved in r ecombinat ion (cr ossover )
at meiosis.
• Maximum r ecombinat ion fr equency possible is 50 %.
• Cr ossing over is absent in case of male Drosophilla and female Bombyx mori (silkwor m)
Interference
Cr ossing over t ake place at chaismat a. These ar e physical st r uct ur es involving t wo chr omat ids.
Not sur pr isingly t he pr esence of one chaismat a in a par t icular chr omosome r egion can r educe
t he fr equency of ot her s for ming close t o it . This is t er med as int er fer ence.
• The ext ent of int er fer ence is calculat ed as coe ffi ci e n t of coi n ci d e n ce (S), which is
equal t o obser ved number of double cr ossover divided by t he expect ed number of
double cr ossover .
• Coeffi ci en t of I n t er fer en ce is 1 minus coff of coincidence (1 – S). If it comes t o be
posit ive values int er fer ence is t er med as posit ive while if t he value is negat ive it is
t er med as negat ive int er fer ence.
II. Dominance
Mendel act ually invest igat ed complet e dominance, i.e. dominant gene t ot ally masks t he r ecessive.
A. Incomplete dominance
A blending of t he effect s of t wo alleles which cr eat es an int er mediat e for m. E.g. RR – r ed,
r r – whit e, Rr – pink. At fir st t his appear s t o be blending inher it ance – but use a Punnet t squar e
t o pr edict t he color s obt ained fr om cr ossing t wo pinks (Rr ).
B. Codominance
Bot h genes ar e expr essed, e.g. human AB blood t ypes.
III. Multiple alleles
Thr ee or mor e alleles for same gene, e.g. t hr ee alleles for human blood gr oups A, B, O wit h O
t he r ecessive and A and B codominant s. Human blood t ypes exhibit bot h codominance and
mult iple alleles. B. Ther e ar e about 100 alleles for hair color in mammals.
IV. Polygenic Inheritance
Tr ait s ar e det er mined by mor e t han one pair of genes. E.g. human eye color and height .
Ther efor e gener ally get cont inuous var iat ion in char act er ist ics and a bell shaped cur ve.
V. Pleiotropy
One allele can affect t wo or mor e t r ait s.E.g. sickle cell anemia is caused by a single mut at ion
in t he hemoglobin gene. It pr oduces a number of effect s.
Genetics 195
VI. Environment may affect the expression of genes. Examples
Eg.1. Siamese cat s have dar k ear s, nose and paws because t hey ar e colder t han r est of t he
body. The enzyme r esponsible for t he dar k color only funct ions at lower t emper at ur es (r ecall
effect of t emper at ur e on enzymes).
Eg.2. Above wat er ver sus submer ged leaves on some plant s ar e mar kedly differ ent .
Eg.3. Ident ical t wins have ident ical genes but exer cise, diet and a number of ot her fact or s
may cause t hem t o have differ ent phenot ypes.
VII. Cytoplasmic inheritance
Recall t hat plast ids and mit ochondr ia have t heir own DNA. Ther efor e some char act er ist ics
may be under t he cont r ol of genes in t he cyt oplasm.
Eg.1. In plants cytoplasm ic inheritance from p l a s t i d s pr oduces t he var iegat ed leaves of
some lea ves, e.g. Coleus.
Eg.2. Male st er ilit y in some cult ivat ed plant s is due t o a mi t och on d r i a l ge n e. This is
useful for pr oducing ar t ificial hybr ids because t he st amens do not have t o be r emoved t o pr event
self-pollinat ion.
VIII. Sex chromosomes
I n ma ny or ga nisms, pa r t icula r ly a nima ls, t he ma les a nd fema les ma y ha ve one pa ir of
chr omosomes t hat ar e differ ent and t hey det er mine t he sex of t he individual. These ar e called
sex chr omosomes.
A. Humans have 22 pair s of aut osomes and one pair of sex chr omosomes. In humans t he
male, Y chr omosome, car r ies a mast er gene called SRY (Sex det er mining Region of t he Y
chr omosome) which causes t he development of male hor mones and r epr oduct ive or gans.
ALLELES AND THEIR INTERACTIONS
• Differ ences in alleles of genes ar e slight differ ences in t he DNA sequence at t he same
locus, which r esult in slight ly differ ent pr ot ein pr oduct s. Some alleles ar e not simply
dominant or r ecessive. Ther e may be many alleles for a single char act er or a single
allele may have mult iple phenot ypic effect s.
New alleles arise by mutation
• Differ ent alleles exist because any gene is subject t o mut at ion, or change, t o a st able,
her it able new for m. Alleles can r andomly mut at e t o become a differ ent allele depending
on DNA sequence changes. Wild type is a t er m used for t he most common allele in
t he populat ion. Ot her alleles, oft en called mut ant alleles, may pr oduce a phenot ype
differ ent fr om t hat of t he wild-t ype allele. A genet ic locus is called polymorphic if t he
wild-t ype allele has a fr equency less t han 99% in a populat ion (t hat is, if mor e t han
1% of t he alleles at t hat locus ar e mut ant alleles).
Many genes have multiple alleles
• A populat ion might have mor e t han t wo alleles for a given gene. The ABO blood t ypes
are an example of multiple alleles. In r abbit s, coat color is det er mined by one gene
wit h four differ ent alleles. Five differ ent color s r esult fr om t he combinat ions of t hese
196 CSIR-NET Life Sciences
alleles. Even if mor e t han t wo alleles exist in a populat ion, any given individual can
have no mor e t han t wo of t hem: one fr om t he mot her and one fr om t he fat her .
Incomplete Dominance-Dominance is usually not complete
• Het er ozygot es may show an int er mediat e phenot ype. For example, r ed-flower ed
snapdr agons (Mir abalis jalapa; 4 O” Clock plant ) when cr ossed wit h whit e will gener at e
pink-flower ed plant s. This phenot ype might seem t o suppor t t he b l e n d i n g t h e or y
(The blending t heor y pr edict s pink F
1
pr ogeny). The F
2
pr ogeny, however , demonst r at e
Mendelian genet ics. When t he F
1
pink individuals self-fer t ilize, t he F
2
pr ogeny have
a phenot ypic r at io of 1 r ed: 2 pink: 1 whit e (The blending t heor y pr edict s all pink F
2
pr ogeny). This mode of inher it ance is called incomplet e dominance.
• The phenot ypic out comes for snapdr agon flower color and incomplet e dominance in
gener al can be explained biochemically. One allele of t he gene codes for an enzyme
t hat funct ions in t he pr oduct ion of t he r ed color . The enzyme coded by t he ot her
allele is not funct ional in pigment pr oduct ion. The r ed-flower ed plant s have t wo
funct ional copies of t he gene and pr oduce enough enzyme t o make r ed flower s. The
pink-flower ed plant s have one funct ional allele, just enough t o cr eat e a pink color .
Neit her allele in whit e-flower ed plant s is capable of making t he funct ional enzyme,
so t he r esult ing flower s ar e whit e.
In co-dominance, both alleles are expressed
• In codominance, t wo differ ent alleles for a gene pr oduce t wo differ ent phenot ypes in
t he het er ozygot es. The AB of t he human ABO blood gr oup syst em is an example. The
alleles for blood t ype ar e I
A
, I
B
and I
O
. They all occupy one locus. These alleles
det er mine which ant igens (pr ot eins) ar e pr esent on t he sur face of r ed blood cells.
• These ant igens r eact wit h pr ot eins called ant ibodies in t he ser um of cer t ain individuals.
The r esult is r ed blood cell agglut inat ion, or clumping, which may be fat al for t hose
individuals.
• Individuals wit h t wo I
A
, or wit h I
A
and I
O
, ar e t ype A. Individuals wit h t wo I
B
, or wit h
I
B
and I
O
, ar e t ype B.
• Individuals wit h I
A
and I
B
ar e t ype AB.
• Individuals wit h I
O
and I
O
ar e t ype O.
• Bot h I
A
and I
B
ar e expr essed when pr esent , and bot h pr oduce an ant igen.
• This is why t hey ar e called codominant .
• I
O
is a recessive trait and is the absence of either the A or B antigen.
Pleiotropy–Some alleles have multiple phenotypic effects
• Pleiot r opy is t he effect of a single gene on mor e t han one char act er ist ic (phenot ype).
An example is t he “fr izzle-t r ait ” in chickens. The pr imar y r esult of t his gene is t he
pr oduct ion of defect ive feat her s. Secondar y r esult s ar e bot h good and bad; good include
incr eased adapt at ion t o war m t emper at ur es, bad include incr eased met abolic r at e,
decr eased egg-laying, changes in hear t , kidney and spleen. Cat s t hat ar e whit e wit h
blue eyes ar e oft en deaf, whit e cat s wit h a blue and an yellow-or ange eye ar e deaf on
t he side wit h t he blue eye. Sickle-cell anemia is a human disease or iginat ing in war m
Genetics 197
low land t r opical ar eas wher e malar ia is common. Sickle-celled individuals suffer
fr om a number of pr oblems, all of which ar e pleiot r opic effect s of t he sickle-cell allele.
Anot her example is t he color at ion pat t er n and cr ossed eyes of Siamese cat s, which
ar e bot h caused by t he same allele. These unr elat ed char act er s ar e caused by t he
same pr ot ein pr oduced by t he same allele.
GENE INTERACTIONS
Some genes (Epistatic) alter the effects of other (Hypostatic) genes
• Epist asis occur s when t he alleles of one gene cover up or alt er t he expr ession of
alleles of anot her gene.
• Re ce s s i ve Ep i s t a s i s (An example is coat color in mice.) Ra t i o 9:3:4.
• The B allele det er mines a banded pat t er n, called agout i. The r ecessive b allele r esult s
in unbanded hair s. The genot ypes BB or Bb ar e agout i. The genot ype bb is solid
color ed (black). Anot her locus det er mines if any color at ion occur s. The genot ypes AA
and Aa have color and aa ar e albino. The aa genot ype blocks all pigment pr oduct ion.
The mice het er ozygous for bot h genes ar e agout i. The F
2
phenot ypic r at io is 9 a gou t i :3
b l a ck :4 wh i t e. The cor r esponding genot ypes ar e 9 agout i (1 BBAA + 2 BbAA + 4
BbAa):3 black (1 bbAA + 2 bbAa):4 albino (1 BBaa + 2 Bbaa + 1 bbaa).
• Comp l e me n t a t i on (9:7): As anot her example of epist asis, imagine t hat when a t r ue-
br eeding whit e-flower ed plant is cr ossed t o a t r ue-br eeding pur ple-flower ed plant ,
t he F
1
ar e all pur ple. When t he pur ple F
1
plant s ar e self-cr ossed, 9 pur ple for ever y
7 whit e ar e obser ved in t he F
2
pr ogeny. This r at io is differ ent fr om what would be
expect ed if pur ple wer e simply dominant t o whit e. The r at io pr ovides a clue t o t he
r elat ionship of t wo differ ent genes (A and B) necessar y t o cr eat e t he pur ple pigment
in t his plant . Suppose t he gene at locus A has t wo alleles: A, which is dominant and
codes for enzyme A, and a, which is r ecessive and codes for a nonfunct ional enzyme.
Suppose also t hat t he gene at locus B has t wo alleles t hat follow t he same pat t er n: B
is dominant and codes for enzyme B, and b is r ecessive and codes for a nonfunct ional
enzyme. The following biochemical pat hway for pr oduct ion of pur ple pigment could
explain t he r at io in t he dihybr id cr oss:
colorless enzyme A colorless enzyme B purple
precursor → intermediate → pigment
• Dominant alleles ar e necessar y at bot h t he A and B loci t o pr oduce pur ple pigment .
Bot h enzyme r eact ions, A and B, must t ake place. Such genes ar e called complement ar y
genes. A Punnet t square for t he dihybrid cross, wit h Xs drawn t hrough t he boxes of
offspring t hat cannot produce pigment , shows clearly t he 9 pu r pl e:7 wh i t e r a t i o.
Some other ratios of Gene Interactions
• Su p p l e me n t a r y ge n e a n d Re ce s s i ve e p i t a s i s : 9:3:4
• Domi n a n t e p i t a s i s : 12:3:1
• Du p li ca t e Ge n e : 15:1
• I n h i b i t or y ge n e : 13:3
• Colla b or a t or ge n e : 9:3:3:1
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Hybrid vigor results from new gene combinations and interactions
• For cent ur ies, it has been known t hat when t wo homozygous st r ains of plant s or
animals ar e cr ossed, t he offspr ing ar e phenot ypically st r onger , lar ger , and mor e
vigor ous t han eit her par ent . In t he ear ly t went iet h cent ur y, G. H. Shull cr ossed t wo
var iet ies of cor n, and t he yield went fr om 20 t o 80 bushels per acr e. This is called
eit her h yb r i d vi gor or h e t e r os i s . Hybr idizat ion is now a common agr icult ur al
pr act ice t o incr ease pr oduct ion in plant s. A hypot hesis called over d omi n a n ce pr oposes
t hat t he het er ozygous condit ion in cer t ain impor t ant genes is super ior t o eit her
homozygot e.
POLYGENIC INHERITANCE
• Polygenic inher it ance is a pat t er n r esponsible for many feat ur es t hat seem simple on
t he sur face. Many t r ait s such as height , shape, weight , color , and met abolic r at e ar e
gover ned by t he cumulat ive effect s of many genes. Polygenic t r ait s ar e not expr essed
as absolut e or discr et e char act er s, as was t he case wit h Mendel’s pea plant t r ait s.
Inst ead, polygenic t r ait s ar e r ecognizable by t heir expr ession as a gr adat ion of small
differ ences (a cont inuous var iat ion). The r esult s for m a bell shaped cur ve, wit h a
mean value and ext r emes in eit her dir ect ion.
• Height in humans is a polygenic t r ait , as is color in wheat ker nels. Height in humans
is not discont inuous. If you line up t he ent ir e class a cont inuum of var iat ion is evident ,
wit h an aver age height and ext r emes in var iat ion (ver y shor t [ver t ically challenged?]
and ver y t all [ver t ically enhanced]). Tr ait s showing cont inuous var iat ion ar e usually
cont r olled by t he addit ive effect s of t wo or mor e separ at e gene pair s. This is an
example of polygenic inher it ance. The inher it ance of each gene follows Mendelian
r ules.
• Usually polygenic t r ait s ar e dist inguished by
1. Tr ait s ar e usually quant ified by measur ement r at her t han count ing.
2. Two or mor e gene pair s cont r ibut e t o t he phenot ype.
3. Phenot ypic expr ession of polygenic t r ait s var ies over a wide r ange.
• Human polygenic t r ait s include: Height , Weight , Eye Color , Int elligence, Skin Color ,
Many for ms of behaviour .
Genetics 199
• Individual her it able char act er s ar e oft en found t o be cont r olled by gr oups of sever al
genes, called polygenes. Each allele int ensifies or diminishes t he phenot ype. Var iat ion
is cont inuous or quant it at ive (“adding up”). Examples of cont inuous char act er s ar e
height , skin color , and possibly int elligence.
THE ENVIRONMENT AFFECTS GENE ACTION
• Genot ype and envir onment int er act t o det er mine t he phenot ype of an or ganism.
Var iables such as light , t emper at ur e, and nut r it ion can affect t he t r anslat ion of
genot ype int o phenot ype. For example, t he dar kness of t he fur on ext r emit ies of a
Siamese cat is affect ed by t he t emper at ur e of t hat r egion. Dar kened ext r emit ies
nor mally have a lower t emper at ur e t han t he r est of t he body. The color at ion can be
manipulat ed exper iment ally. The pr opor t ion of individuals in a gr oup wit h a given
genot ype t hat expr ess t he cor r esponding phenot ype can somet imes be measur ed,
and t he measur e is called p en et r a n ce.
• The exp r essi vi t y of t he genot ype is t he degr ee t o which it is expr essed in an individual.
• An example is hereditary hemochromatosis. This disease causes abnormally high levels
of iron t o accumulat e in t he liver and ot her organs of affect ed people. S ome people
wit h t he disease accumulat e t oxic levels of iron, and ot hers accumulat e levels just
above normal.
• The influence of envir onment on genot ype or phenot ype can be st udied wit h ident ical
t wins, especially when t hey ar e separ at ed fr om bir t h and r ear ed apar t in subst ant ially
differ ent envir onment s.
GENES AND CHROMOSOMES
• How do we det er mine t he or der and dist ance bet ween t he genes t hat ar e locat ed on
t he same chr omosome? A syst em was fir st developed in Thomas Hunt Mor gan’s fly
lab in 1909. The biological model used was t he fr uit fly, Drosophila melanogaster,
which is st ill used t oday in chr omosomal st udies.
Linked genes are on the same chromosome
• Mendel’s second law, independent assor t ment , failed t o be univer sal. One ear ly
except ion was found in Drosophila when cr ossing flies t hat wer e hybr ids for t wo
par t icular alleles (body color and wing size) wit h flies t hat wer e r ecessive for bot h
alleles (a t est cr oss). The r esult s wer e not t he expect ed 1:1:1:1, but inst ead, t wo of
t he genot ypes occur r ed at a fr equency higher t han t he ot her t wo. These r esult s
make sense if t he t wo loci ar e on t he same chr omosome, and t hus t heir inher it ance
is linked. All t he loci on a given chr omosome make up a linkage gr oup. Absolut e or
t ot al linkage of all loci is however , ext r emely r ar e.
• Dr osoph ila h ave j u st 3 pai rs of au t osomal ch romosomes an d on e pai r of sex
chromosomes. The normal fly is diploid and has 8 chromosomes. The fr uit fly has
t housands of genes on just 4 pair s of chr omosomes. Ther efor e, many exist t oget her
on t he sa me chr omosome.
200 CSIR-NET Life Sciences
Genes can be exchanged between chromatids
• When t wo homologous chr omosomes physically exchange cor r esponding segment s
dur ing pr ophase I of meiosis, genet icist s call it cr ossing over .
• Recombinat ions occur at ch i a s ma t a d u r i n g p a ch yt e n e of me i os i s -I . If just a few
exchanges occur , genes t hat ar e closer t oget her t end t o st ay t oget her . The far t her
apar t on t he same chr omosome genes ar e, t he mor e likely t hey will separ at e dur ing
r ecombina t ion. The t wo ext r emes a r e independent a ssor t ment a nd complet e or
absolut e linkage.
• The pr ogeny r esult ing fr om cr ossing over appear in r epeat able pr opor t ions, called
t h e r e comb i n a n t fr e q u e n cy. Gr eat er r ecombinat ion fr equencies ar e obser ved for
genes t hat ar e far t her apar t on t he chr omosomes because a chiasma is mor e likely t o
cut bet ween genes t hat ar e far apar t t han genes t hat ar e closer t oget her .
Geneticists make maps of eukaryotic chromosomes
• Alfr ed St ur t evant , an under gr aduat e st udent wor king in Mor gan’s fly r oom, r esolved
t he puzzling quest ion of t he deviat ion of r esult s fr om t he expect ed r at ios by suggest ing
t hat as t he dist ance bet ween t wo genes on a chr omosome incr eases, so does t he
likelihood t hat t hey will separ at e and r ecombine. The closer loci ar e on a chr omosome,
t he less likely t hey will separ at e and r ecombine in meiosis.
• A map unit is a r ecombinat ion fr equency of 0.01 (or a 1% r ecombinat ion). It is r efer r ed
t o as a ce n t i mor ga n (cM).
EUKARYOTIC CHROMOSOMES
• Apar t fr om gamet es, most human cells cont ain t wo full set s of genet ic infor mat ion,
one fr om t he mot her and t he ot her fr om t he fat her . Eukar yot es have mor e t han one
chr omosome, and t he number var ies fr om or ganism t o or ganism; for example, humans
have 46 and hor ses have 64. Each eukar yot ic chr omosome consist s of a single, double-
st r anded molecule of DNA; t he molecule is ext r emely long r elat ive t o t he size of t he
cell. Many pr ot eins associat e wit h t he DNA molecule. Aft er t he DNA of a chr omosome
r eplicat es dur ing S phase, each chr omosome consist s of t wo joined chr omat ids. The
t wo chr omat ids ar e joined at a r egion called t he cent r omer e.
Chromatin consists of DNA and proteins
• The complex of t he DNA and pr ot eins is called chr omat in. The DNA car r ies t he
genet ic infor mat ion. The pr ot eins or ganize t he DNA physically and r egulat e t he
act ivit ies of t he DNA. By mass, chr omat in consist s of about equal par t s DNA and
pr ot ein. Chr omat in condenses, becoming highly coiled and compact ed dur ing mit osis
or meiosis. This makes it possible t o get copies of t he DNA int o separ at e cells. Wit hout
condensat ion of t he chr omat in, t he st r ands, which ar e much longer t han t he cell,
would fail t o pr oper ly par t it ion int o t he t wo sist er cells.
Chromatin proteins organize the DNA in chromosomes
• DNA of a human cell has a t ot al lengt h of 2 met er s. The nucleus is just 5 mm in
diamet er . Dur ing int er phase, t he DNA is decondensed or “unwound,” alt hough pr ot eins
Genetics 201
st ill package it . Int er phase chr omosomes ar e wr apped ar ound pr ot eins called hist ones.
These wr aps of DNA and hist one pr ot eins ar e called n u cl e os ome s (11nm diamet er )
and r esemble beads on a st r ing.
• Ther e ar e five classes of hist ones. The cor e of a nucleosome cont ains eight hist one
molecules, t wo each fr om four of t he hist one classes. Ther e ar e 146 base pair s of
DNA wr apped ar ound t he cor e, or 1.65 t ur ns of DNA. One molecule fr om t he r emaining
hist one class, hist one H1, clamps t he DNA t o t he cor e, and helps for m t he next level
of packaging.
• Dur ing mit osis and meiosis, t he chr omat in becomes even mor e coiled and condensed.
Telomerase, Aging and Cancer
• The ends of chr omosomes ar e called t elomer es. When DNA is r eplicat ed t he t elomer es
ar e oft en not duplicat ed pr oper ly and t he chr omosome becomes a lit t le shor t er aft er
each r eplicat ion.
• Some scient ist s believe t hat t he gr adual shor t ening of t he chr omosomes causes cell
aging and event ual deat h (most cells in t he body can duplicat e only 50-100 t imes
befor e t hey die). Cells which divide oft en (ger m cells, st em cells and cancer cells)
have high levels of an enzyme called t elomer ase which allows t he t elomer es t o be
duplicat ed pr oper ly.
• Telomer ase may make t he cells pot ent ially immor t al. Inhibit or s of t elomer ase may
someday be useful in cancer t her apy.
• Met hot r exa t e, a minopt er in a nd a minopt hr in a r e a nt ica ncer ous dr ug which a r e
nucleot ide analog and inhibit t he act ivit y of enyme involved in salvage pat hway of
nucleot ide biosynt hesis.
GENE EXPRESSION
• Two st eps ar e used t o expr ess a gene:
• Tr anscr ipt ion makes a single-st r anded RNA copy of a segment of t he DNA.
• Tr anslat ion uses infor mat ion encoded in a por t ion of t he RNA t o make a polypept ide.
• In eukaryot es, t hese t wo st eps are physically separat ed in t ime and space.
RNA differs from DNA
• RNA is single-st r anded. The sugar in RNA is r ibose, not deoxyr ibose since 2’OH in
r ibose was act ing as nucleophile and was involved in Aut o-cat alyt ic act ivit y t o make
genet ic mat er ial st able 2’ Oxygen was r emoved t o yield deoxyr ibose. Wher ever t hymine
is found in DNA, it is r eplaced by ur acil in RNA. Dur ing evolut ion ur acil of RNA was
changed int o Thymine of DNA so as t o help r epair machiner y t o r ecognize deaminat ed
cyt osine. RNA can fold over and base-pair wit h it self.
Information flows in one direction when genes are expressed
• Fr ancis Cr ick’s cent r al dogma st at ed t hat DNA codes for RNA, and RNA codes for
pr ot ein; t hat is, once infor mat ion passes int o pr ot ein, it cannot get out again. Messenger
RNA (mRNA) moves fr om t he nucleus of eukar yot ic cells int o t he cyt oplasm wher e it
202 CSIR-NET Life Sciences
ser ves as a t emplat e for pr ot ein synt hesis. Tr ansfer RNA (t RNA) is t he link bet ween
t he code of t he mRNA and t he amino acids of t he polypept ide. The t RNA molecules
specify t he cor r ect amino acid.
RNA viruses modify the central dogma
• RNA vir uses a r e vir uses t ha t use RNA a s t heir infor ma t ion molecule dur ing
t r ansmission. Examples ar e t he influenza vir us and poliovir us. HIV and cer t ain
t umor vir uses (Re t e r ovi r u s e s ) have RNA as t heir infect ious infor mat ion molecule,
but conver t it t o a DNA copy inside t he host cell, t hen use it t o make mor e RNA wit h
t he help of r ever se t r anscr ipt ase (RNA d e p e n d e n t DNA p ol yme r a s e) by pr ocess
known as r ever se t r anscr ipt ion.
Transcription: DNA-Directed RNA Synthesis
• RNA polymer ase is t he enzyme t hat uses DNA as a t emplat e t o make RNA in dir ect ion
5’®3’. J ust one of t he st r ands of a gene’s DNA is used t o make t he RNA. This st r and
is ca lled t he t empl at e st rand (3’®5’) a nd is used for t r a nscr ipt ion. The ot her
unt r anscr ibed st r and is called t he complement ary st rand (sense st rand/ non-coding
strand). For differ ent genes in t he same DNA molecule, however , t he r oles of t hese
st r ands may be r ever sed.
• The cont inuous double helix of DNA ha s ma ny r egions t ha t a r e r ea d by RNA
polymer ase. The DNA double helix par t ly unwinds t o ser ve as t emplat e. As t he RNA
t r anscr ipt for ms, it peels away, allowing t he alr eady t r anscr ibed DNA t o be r ewound
int o t he double helix.
Initiation of transcription requires a promoter and an RNA polymerase
• Tr anscr ipt ion of a gene begins at a pr omot er , which is a cer t ain sequence of DNA.
Ther e is at least one pr omot er for each gene t o be t r anscr ibed int o mRNA. The RNA
polymer ase binds t o t he pr omot er r egion when condit ions allow. The pr omot er
sequence dir ect s t he RNA polymer ase as t o which of t he double st r ands is t he t emplat e
and in what dir ect ion t he RNA polymer ase should move. In effect , t he pr omot er s
ser ve as punct uat ion mar ks for t he t r anscr ipt ion pr ocess. RNA is synt hesized in t he
5' -t o-3' direct ion, moving along t he t em pl a t e DNA i n t h e 3' -t o-5' d i r ect i on .
• In pr oka r yot es pr omot er s a r e cent er ed a t –10 (TATAAT pr ibnow box) a nd –35
(TTGACA). In eukar yot es 80% of genes have a TATA box cent er ed at –25. Fur t her
upst r eam, wit hin 100-200 ar e boxes or element s—The CAAT box, GC box and eight
base pair oct amer box. Not all pr omot er s ar e ident ical.
• Some bind RNA polymer ase mor e effect ively; t his causes t hem t o be t r anscr ibed
mor e fr equent ly, when ot her condit ions allow. For cor r ect init iat ion (s) fact or is
r equir ed which is r elea sed a ft er init ia t ion. Pr oka r yot es ha ve one t ype of RNA
polymer ase t hat t r anscr ibes mRNA, t RNA, and r RNA.
• In E. Coli t he RNA cor e enzyme has four subunit a
2
(160 KD each) b(150 KD) b’(155KD)
and s (70 KD)fact or at t ach for init iat ion t o for m holoenzyme. b -subunit is involved in
phosphodiest er bond for mat ion (c a t a l yt i c s i t e), bot h b and b’ subunit s par t icipat e in
DNA binding and a subunit have sever al act ivit ies including r ecognit ion of UP element s
in pr omot er .
Genetics 203
• Ant ibiot ic r i fa myci n inhibit s pr okar yot ic RNA polymer ase by act ing at t he cat alyt ic
site (b s u b u n i t ). Anot her ant ibiot ic known as s t r e p t ol yd i gi n , blocks t he RNA chain
elongat ion.
• Eukar yot es have t hr ee differ ent RNA polymer ases which can be separ at ed by i on
e xch a n ge ch r oma t ogr a p h y: RNA polymer ase I (found in nucleolus, t r anscr ibes
r RNA), II (mRNA), and III (t -RNA; bot h II and III pr esent in nucleoplasm).
• RNA polymer a se II ma kes a ll mRNA in euka r yot es a nd is highly sensit ive t o
a-a ma n i t i n obt ained fr om mushr ooms of genus Ama n i t a . RNA polymer ase I is
insensit ive while RNA pol III shows sensit ivit y at high dose.
• r RNA synt hesis ca n be inhibit ed by a c t i n o my c i n D a s it select ively in ser t s
(int er calcat es) in bet ween GC-r ich r egion of DNA and since r RNA genes ar e r ich in
GC sequences.
RNA polymerase elongates the transcript
• Aft er binding, RNA polymer ase unwinds t he DNA about 20 base pair s at a t ime and
r eads t he t emplat e in t he 3' -t o-5' dir ect ion. The RNA t r anscr ipt is ant ipar allel t o t he
DNA t emplat e st r and. Ener gy for synt hesis comes fr om t he r emoval and br eakdown
of t he pyr ophosphat e gr oup fr om each nucleoside added. Tr anscr ipt ion er r or s for
RNA polymer ases ar e high r elat ive t o DNA polymer ases; a mist ake occur s for ever y
10
4
t o 10
5
bases incor por at ed. These ar e er r or s in t he copies, however , not t he or iginal
DNA mast er , so t hey ar e less likely t o be har mful.
• Tr anscr ipt ion fact or s ar e r equir ed for RNA polymer ase t o r ecognize pr omot er s. TFIID
is t he basal t r anscr ipt ion fact or in eukar yot es which int er act dir ect ly wit h TATA box
wit h help of Tat a Binding Pr ot ein (TBP) and eight or mor e ot her TAFs (TBP associat ed
fact or s).
• The most common mot if for pr ot eins int er act ing pr omot er s in pr okar yot es is Heli x
t u r n h e l i x while in eukar yot es ar e le u ci n e Zi p p e r a n d Zi n c fi n ge r (e g. TF I I D,
TF I I I A) pr ot eins. Home od oma i n s pr ot eins coded by Homeot ic genes ar e highly
conser ved being similar in st r uct ur es in insect Dr osophila and in ver t ebr at es, including
humans.
Transcription terminates at particular base sequences
• Par t icular base sequences in t he DNA specify t er minat ion (Gener ally GC r ich, for ms
hair pin loop in mRNA followed by A r ich sequence). Gene mechanisms for t er minat ion
var y. For some genes, t he newly for med t r anscr ipt simply falls away fr om t he DNA
t emplat e and t he RNA polymer ase. For ot her genes, a helper pr ot ein (Rh o fa ct or )
pulls t he t r anscr ipt away by it s unwinding (h e l i ca s e) act ivit y.
• In pr okar yot es, t r anslat ion of t he mRNA oft en begins befor e t r anscr ipt ion is complet e.
• In eukar yot es, t he pr ocess is mor e complicat ed and involves a spat ial separ at ion as
well as fur t her pr ocessing.
THE GENETIC CODE
•DNA codes for RNA by the transcription process. mRNA is read in three-base contiguous segments
called codons.
204 CSIR-NET Life Sciences
•The number of different codons possible is 64, because each position in the codon can be occupied by one
of four different bases. Four possibilities for the first base, times four for the second, times four for the third
yields 64 possibilities. The 64 possible codons code for only 20 amino acids and the start and stop signals
found in all mRNA molecules. AUG, which codes for methionine, is called the start codon. The start codon
initiates translation. Three of the possible codons are stop codons (UAA, UAG, and UGA). Stop codons
direct the ribosome to stop reading the mRNA since there is no t-RNA for them; that is, they end translation.
The genetic code is redundant but not ambiguous
• Aft er subt r act ing st ar t (codes for met hionine) and st op codons, t he r emaining 60
codons code for 19 differ ent amino acids. This means t hat many amino acids have
mor e t han one codon. Thus t he code is r edundant . But t he code is not ambiguous.
Each codon is assigned only one amino acid, not t wo or t hr ee possible amino acids.
• The t RNA mol ecul es t hat have t he correct ami no aci ds at t ached det ermi ne t he
assignment of amino acids t o t he mRNA codons. The genet ic code is near ly univer sal,
applying t o all species on our planet . Minor var iat ions ar e found wit hin mit ochondr ia
and chlor oplast s; ot her except ions ar e few and slight . This common genet ic code has
gr eat implicat ions in genet ic engineer ing.
• In mit ochondr ia of Dr osophila, yeast , higher plant s UGA is codon for t r ypt ophan
r at her t hen st op. In mammalian mit ochondr ia including human, AGG and AGA, t hey
ar e st op codon inst ead of ar ginine.
Biologists broke the genetic code by using artificial messengers
• Decoding br eakt hr oughs st ar t ed in 1961. Ear ly on, t he likelihood of a t hr ee-let t er
codon was post ulat ed. Nir enber g pr epar ed an ar t ificial mRNA in which all bases wer e
ur acil. Incubat ed wit h r equir ed addit ional component s, t he poly U mRNA led t o
synt hesis of a polypept ide chain consist ing only of phenylalanine amino acids. UUU
appear ed t o be t he codon for phenylalanine. Ot her codons wer e decipher ed fr om t his
st ar t ing point .
• An addit ional t echnique finished t he decipher ing. Simple synt het ic mRNA’s, t hr ee
nucleot ides long, could bind t o r ibosomes. This complex t hen caused t he t RNA-amino
acid t o bind accor ding t o t he t hr ee-let t er codon. Using t his t echnique, t he code was
fully decipher ed.
• Radioactive labeling was also used to decipher the code.
Preparation for Translation: Linking RNA’s, Amino Acids, and Ribosomes
• Tr anslat ion occur s at r ibosomes, which ar e molecular pr ot ein synt hesizing machines
t hat hold mRNA and t RNA in place. In pr okar yot es, r ibosomes bind t o mRNA as it is
being synt hesized. In eukar yot es, mRNA is made in t he nucleus, and t r anslat ion
occur s at t he r ibosomes in t he cyt oplasm.
• To assur e pr ot ein specificit y: The t RNA’s must r ead mRNA cor r ect ly & it must car r y
t he cor r ect amino acids.
Transfer RNA’s carry specific amino acids and bind to specific codons
• Specific t RNA molecules funct ion as adapt er s. Each car r ies an amino acid, associat es
wit h mRNA molecules, and int er act s wit h r ibosomes. A t RNA molecule has 75 t o 80
Genetics 205
nucleot ides. It has a t hr ee-dimensional shape maint ained by complement ar y base
pair ing and hydr ogen bonding. At t he 3' end of ever y t RNA molecule is a sit e t o which
it s specific amino acid binds covalent ly. Midpoint in t he sequence ar e t hr ee bases
called t he ant icodon.
• The ant icodon is t he cont act point bet ween t he t RNA and t he mRNA. The ant icodon
is complement ar y (and ant ipar allel) t o t he mRNA codon. The codon and ant icodon
unit e by complement ar y base pair ing.Ther e ar e fewer ant icodon codes t han mRNA
codons. This is possible beca use some codon-a nt icodon int er a ct ions t oler a t e a
mismat ch at t he 3' base of t he mRNA. This is called wob b le, but does not allow t he
genet ic code t o be ambiguous.
• The t hr ee-dimensional shape of t he t RNA’s allows t hem t o combine wit h t he binding
sit es of t he r ibosome.
Activating enzymes link the right tRNA’s and amino acids
• The cor r ect amino acids ar e at t ached t o t he cor r ect t RNA’s by a family of act ivat ing
enzymes called aminoacyl-t RNA synt het ases. Each act ivat ing enzyme is specific for
one amino acid and it s t RNA. The enzyme has a t hr ee-par t act ive sit e t hat binds a
specific amino acid, ATP, and a specific t RNA, which is char ged wit h a high-ener gy
bond. This bond pr ovides t he ener gy for making t he pept ide band t hat will join adjacent
amino acids.
• The r eact ions have t wo st eps: Enzyme + ATP + AA ® enzyme—AMP—AA + PP
i
Enzyme—AMP—AA + tRNA → enzyme + AMP + tRNA—AA
The ribosome is the staging area for translation
• Each r ibosome has t wo subunit s, a lar ger one and a smaller one. The lar ge one in
eukar yot es has t hr ee differ ent associat ed r RNA molecules and 45 differ ent pr ot eins.
The smaller subunit has one r RNA and 33 differ ent pr ot ein molecules. When t hey
ar e not t r anslat ing, t he t wo subunit s ar e separ at e. Ribosomes of pr okar yot es ar e
somewha t sma ller , a nd t heir r ibosoma l pr ot eins a nd r RNA’s a r e differ ent . The
differ ent pr ot eins and r RNA’s ar e held t oget her by ionic bonds and hydr ophobic for ces,
not covalent bonds. The st r uct ur e can self-assemble if disassembled by det er gent s.
• Ribosomes ar e simply molecular fact or ies and ar e nonspecific. They combine wit h
any mRNA and all t RNA’s. The lar ge subunit has four sit es wher e t RNA molecules
bind. The T si t e is wher e t he t RNA fir st lands. It is br ought t o t he sit e by a special
pr ot ein, t he T, or t r ansfer fact or . Th e A s i t e is wher e t he t RNA ant icodon binds t o
t he mRNA codon. Th e P s i t e is wher e t he t RNA adds it s amino acid t o t he gr owing
polypept ide chain. The E (e xi t ) s i t e is wher e t he t RNA, less it s amino acid, goes
befor e leaving t he r ibosome.
TRANSLATION: RNA-Directed Polypeptide Synthesis
Translation begins with an initiation complex
• At t he beginning of t r anslat ion, m-RNA at t aches it self t o small subunit of r ibosome
(by s h i n e d a l ga r n o s e q u e n c e complemen t a r y t o 16 S r RNA of r ibosome in
pr okar yot es or at 5' cap in eukar yot es) t o for m an init iat ion complex. The init iat ion
complex includes t he fir st t RNA and it s amino acid, a small subunit of t he r ibosome,
206 CSIR-NET Life Sciences
and an mRNA molecule. This complex is bound t o a r egion upst r eam (t owar d t he 5'
end) of wher e t he act ual r eading of t he mRNA begins. The st ar t codon (AUG) for
met hionine (for myl met hionine in pr okar yot es) designat es t he fir st amino acid in all
pr ot eins. (However , some pr ot eins ar e t r immed aft er synt hesis, and t he met hionine
is t her eby r emoved). In euka r yot es init ia t or codon AUG lies wit hin consensus
sequence known as Kozak sequence. The lar ge subunit t hen joins t he complex. The
pr ocess is dir ect ed by pr ot eins called initiation factors, which use GTP as an ener gy
sour ce.
The polypeptide elongates from the N terminus
• Ribosomes move in t he 5' -t o-3' dir ect ion on t he mRNA. They synt hesize t he pept ide in
t he N t erminus-t o-C t erminus direct ion. The lar ge subunit cat alyzes t wo r eact ions:
Br eakage of t he bond bet ween t he t RNA in t he P sit e and it s amino acid (on t he
polypept ide) and Pept ide bond for mat ion bet ween t his (t RNA-at t ached) amino acid
and t he t RNA in t he A sit e. This is called p e p t i d yl t r a n s fe r a s e act ivit y (by 23 S
r RNA in pr okar yot es). One of t he r RNA’s in t he lar ge subunit s appear s t o par t icipat e
in the catalysis of this reaction. Here we see rRNA acting as the catalyst, or r i boz yme .
Elongation continues and the polypeptide grows
• The fir st t RNA r eleases met hionine, dissociat es fr om t he r ibosome, and r et ur ns t o
t he cyt osol. The second t RNA t hen moves t o t he P sit e. The next char ged t RNA
ent er s t he open A sit e. The pept ide chain is t hen t r ansfer r ed t o t he P sit e. These
st eps ar e assist ed by pr ot eins called elongat ion fact or s.
A release factor terminates translation
• When a st op codon—UAA, UAG, or UGA—ent er s t he A sit e, a r elease fact or and a
wat er molecule ent er t he A sit e, inst ead of an amino acid. The newly complet ed
pr ot ein t hen separ at es fr om t he r ibosome.
REGULATION OF GENE EXPRESSION
• Hi s t on e a ce t yl a t i on occur s bot h in cyt oplasm and nucleus. Cyt oplasmic acet ylat ion
is ca r r ied out by HAT-B (Hist one a cet yl t r a nsfer a se) a nd pr epa r es hist one for
incor por at ion int o nucleosomes. The acet yl gr oups ar e lat er r emoved in nucleus.
Nuclear acet ylat ion of is cat alyzed by HAT-A a nd cor r ela t es wit h t r a nscr ipt ion
act ivat ion. A var iet y of co-act ivat or s have HAT-A act ivit y, which may allow t hen t o
loosen t he associat ion of nucleosides wit h a genes cont r ol r egion, t her eby en h a n ci n g
t r a n scr i pt i on .
• The t r anscr ipt of appr oximat ely 1 in 20 genes ar e subject t o alt er nat e splicing. This
can have pr ofound effect on t he pr ot ein pr oduct of gene. For eg., it can make differ ence
bet ween secr et ed and membr ane bound pr ot ein.
• 5' Ca p of m-RNA ar e made in st eps: Fir st a p h os p h oh yd r ol a s e r emoves t he t er minal
phosphat e; next a gu a n yl t r a n s fe r a s e adds t he capping GMP (fr om GTP). Next , t wo
met h yl t r a n sfer a se met h yla t es N
7
of gu a n os i n e a n d 2' -O-met hyl gr oup of t he
penult imat e nucleot ide. These event s occur ear ly in t r anscr ipt ion pr ocess, befor e t he
chain lengt h r eaches 30 nt . The cap pr ovides pr ot ect ion of t he mRNA fr om degr adat ion,
Genetics 207
enhancement of t he mRNA’s st abilit y; t r anspor t of mRNA t o nucleus and pr oper
splicing of m-RNA.
• P ol ya d e n yl a t i on : Most eukar yot ic mRNA and t heir pr ecur sor s have a chan of AMP
r esidues about 250 nt long at t heir 3' ends. This poly(A) is added post -t r anscr ipt ionally
by p ol y(A) P ol yme r a s e . Poly(A) enhances bot h life t ime and t r anslat ion abilit y of
mRNA; by h el pi n g t o r ecr u i t mRNA t o pol ys omes . An effi ci en t ma mma l i a n
p o l y a d e n y l a t i o n s i g n a l con sist of AAUAAA mot if a bout 20 nt upst r ea m of
polyadenylat ion sit e in pr e-mRNA.
• Gr ou p I i n t r on s such as Tet rahymena 23 S r RNA pr ecur sor , can be r emoved wit hout
pr ot eins (Ri b ozyme).
• Gr ou p I I (GU-AG) int r ons of eukar yot es self splice using an A-br anched lar iat and
spliceosome.
Some antibiotics work by inhibiting translation
• Ant ibiot ics ar e defense molecules. They ar e pr oduced by some fungi and bact er ia.
They have been used t o combat human bact er ial infect ious disease. Ant ibiot ics must
specifically dest r oy micr obial invader s, but not har m t he human host . Some ant ibiot ics
wor k by blocking t he synt hesis of t he bact er ial cell walls, ot her s by inhibit ing pr ot ein
synt hesis.
• In prokaryot es, s t r e p t omyci n affects initiation, k i r r omyci n prevents EF-Tu release,
and e r y t h r o my c i n an d c h l o r a mp h e n i c o l i nhi bi t s t ransl ocase and pept i dyl
t ransferase respect ively (This is t rue for mit ochondrial ribosomes t oo). F u s i d i c a ci d
inhibits translocation by blocking EF-G-GDP release. Because of differ ences bet ween
pr okar yot ic and eukar yot ic r ibosomes, t he human r ibosomes ar e unaffect ed.
• P u r omyci n r esembles an amino acyl t -RNA and so bind t o A sit e, couple wit h t he
pept ide in t he P sit e, and cause pr emat ur e t er minat ion of pr ot ein synt hesis by
r eleasing pept idyl pur omycin.
• Di p t h e r i a t oxi n inhibits EF2 the eukaryote translocase (Equals EF-G in bacteria).
Ri ci n , t he t oxin of cast or bean, is an N-glycosidase t hat removes a single adenine base
from one of t he eukaryot e ribosomal RNA and inact ivat es large subunit . One molecule
of ricin can dest roy a cell cont aining t ens of t housands of ribosomes.
Polysome formation increases the rate of protein synthesis
• Polysomes ar e mRNA molecules wit h mor e t han one r ibosome at t ached (gener ally
ar ound 5). These make pr ot ein mor e r apidly, pr oducing mult iple copies of pr ot ein
simult aneously. They ar e mor e common in pr okar yot es.
Posttranslational Events
• Some pr ot eins r equir e addit ional modificat ion aft er synt hesis befor e t hey become
funct ional. New chemical gr oups might be added t o t he pr ot ein, it might be folded
(wit h t he assist ance of ot her pr ot eins), or it might get t r immed. Most of such mechanism
occur s in golgi complex and t o some ext ent in lumen of ER.
208 CSIR-NET Life Sciences
Chemical signals in proteins direct them to their cellular destinations
• As t he polypept ide chain for ms, it spont aneously folds int o it s t hr ee-dimensional shape.
The amino acid sequence also cont ains an “addr ess label” indicat ing wher e in t he cell
t he polypept ide belongs. All pr ot ein synt hesis begins on fr ee r ibosomes in t he cyt oplasm.
In eukar yot es, as t he pept ide chain is made, infor mat ion on t he nascent por t ion gives
one of t wo set s of inst r uct ions: 1. Finish t r anslat ion and be r eleased t o t he cyt oplasm
and 2. St all t r anslat ion, go t o t he endoplasmic r et iculum, and finish synt hesis at t he
ER sur face.
• Those dest ined t o finish synt hesis in t he cyt oplasm may cont ain infor mat ion in t heir
amino acid sequence t hat specifies wher e t hey belong: t he nucleus, mit ochondr ia, or
per oxisomes.Some of t he pr ot eins t hat ar e t r anspor t ed t o a dest inat ion r equir e
chaper onin pr ot eins and docking pr ot eins at t he membr ane t hat t he pr ot ein must
cr oss t o it s or ganelle dest inat ion.
• Those dest ined for t he ER gener at e an appr oximat ely 25-amino-acid-long hydr ophobic
leader sequence t hat signals t o a signal r ecognit ion par t icle, which is composed of
pr ot ein and RNA. The associat ion of t he signal t o t he signal r ecept or par t icle st alls
any addit ional t r anslat ion. This st all cont inues unt il t he r ibosome at t aches t o a specific
r ecept or pr ot ein on t he sur face of t he ER. Tr anslat ion cont inues wit h t he pr ot ein
moving t hr ough a por e in t he ER membr ane. Some pr ot eins have signals t hat dir ect
t he embedding of t he pr ot ein int o t he ER membr ane. This is when membr ane pr ot eins
of t he ER, Golgi appar at us, lysosomes, and plasma membr ane get posit ioned. Ot her
signals dir ect t he pr ot ein t o t he Golgi appar at us, lysosomes, or t o t he out side of t he
cell. Pr ot eins wit h no signals fr om t he ER go t hr ough t he Golgi appar at us and ar e
secr et ed fr om t he cell.
Many proteins are modified after translation
• It is t he except ion, not t he r ule, t hat t he finished pr ot ein is ident ical t o t he t r anslat ion
fr om t he mRNA code. Modificat ions ar e oft en essent ial t o t he final funct ioning of t he
pr ot ein.
• Pr ot eolysis is t he cleavage of t he pr ot ein t o make a shor t ened finished pr ot ein. Insulin
is an example of a prot ein t hat get s t rimmed. The signal t o go t o t he ER is oft en
cleaved aft er t he pr ot ein get s t her e. The vir us HIV needs a pr ot ease t o cleave a
pr ot ein. One t r eat ment for HIV inhibit s t his enzyme.
• Glycosylat ion involves t he addit ion of sugar s t o t he pr ot ein. Signals in t he amino acid
sequence of t he pr ot ein dir ect t he addit ion of t he sugar s (N-l i n k e d ) in t he ER by
r esident enzymes. Addit ional modificat ions occur in bot h t he Golgi appar at us (O-
li n k e d ) and t he ER.
• Phosphor ylat ion is t he addit ion of phosphat e gr oups t o cer t ain pr ot eins. These addit ions
may be t empor ar y and affect t he act ivit y of t he pr ot ein by changing t he t hr ee-
dimensional shape.
Degradation of proteins
• Select ive dest r uct ion of pr ot eins involves u b i q u i t i n , a small pr ot ein found univer sally
in eukar yot es (but not in pr okar yot es). By an ATP dependent r eact ion it s –COOH
t er minal gr oup becomes linked e-NH
2
of t he side chain of lysine r esidues of it s t ar get
Genetics 209
pr ot ein and once at t ached, t hat pr ot ein is mar ked for dest r uct ion. P r ot e a s ome present
in cyt osol degr ades t he ubiquit in t agged pr ot ein. Ch a p e r on s also appear s t o play a
r ole in t ar get ing pr ot ein in dest r uct ion.
• Lys os ome s may par t icipat e in t he dest r uct ion of longer lived st r uct ur al pr ot ein.
They ar e also r esponsible for t he aut olyt ic dest r uct ion of cell dur ing t he development .
The disappear ance of t adpole’s t ail is example of t his.
SEX DETERMINATION AND SEX-LINKED INHERITANCE
• Somet imes par ent al or igin of a chr omosome does mat t er . Recipr ocal cr osses give
ident ical r esult s when or ganisms ar e diploid. Many organisms have homologous pairs
of all chromosomes except for t hose t hat det ermine sex. The homologous pairs are
called aut osomes; t he unpaired X and Y chromosomes are called sex chromosomes.
Dosage Compensation by X-inactivation in female mammals
How does an or ganism compensat e for t he fact t hat some individuals have a double dosage of
sex-linked genes while ot her s have only one? In female mammals, most diploid cells have only
one fully funct ional X chr omosome.
• The explanat ion for t his pr ocess is known as t he Lyon hypot hesis, pr oposed by t he
Br it ish genet icist Mar y F. Lyon. In females, each of t he embr yonic cells inact ivat es
one of t he t wo X chr omosomes. The inact ive X chr omosome cont r act s int o a dense
object called a Bar r body.
Barr body
Locat ed inside t he nuclear envelope, it is a densely st aining object t hat is an inact ivat ed X
chr omosome in female mammalian cells. Most Bar r body genes ar e not expr essed. They ar e
r eact ivat ed in gonadal cells t hat under go meiosis t o for m gamet es. Female mammals ar e a
mosaic of t wo t ypes of cells, t hose wit h an act ive mat er nal X and t hose wit h an act ive pat er nal
X. Which of t he t wo Xs will be inact ivat ed is det er mined r andomly in embr yonic cells. Aft er an
X is inact ivat ed, all mit ot ic descendant s will have t he same inact ive X. As a consequence, if a
female is het er ozygous for a sex-linked t r ait , about half of her cells will expr ess one allele and
t he ot her cells well expr ess t he alt er nat e allele. Examples of t his t ype of mosaicism ar e color at ion
in calico cat s and nor mal sweat gland development in humans. A woman who is het er ozygous
for t his t r ait has pat ches of nor mal skin and pat ches of skin lacking sweat glands.
• X chr omosome inact ivat ion is associat ed wit h DNA me t h yl a t i on . Me t h yl gr ou p s
(-CH
3
) a t t a ch t o cyt os i n e, one of DNA’s nit r ogenous bases. Bar r bodies ar e highly
met hylat ed compar ed t o act ively t r anscr ibed DNA. What det er mines which of t he
t wo X chr omosomes will be met hylat ed? A r ecent ly discover ed gene, XI ST is act ive
only on t he Bar r body. The pr oduct of t he XIST gene, X-inact ive specific t r anscr ipt , is
an RNA; mult iple copies of XIST at t ach t o t he X chr omosome inact ivat ing it .
Sex is determined in different ways in different species
• In cor n, ever y diploid adult has bot h male and female r epr oduct ive st r uct ur es. The
same is t r ue for peas. This t ype of or ganism is called mon oe ci ou s (“one house”).
Ot her plant s and animals, which have individuals t hat ar e one or t he ot her sex, ar e
called d i oe ci ou s (“t wo houses”).
210 CSIR-NET Life Sciences
• In most dioecious or ganisms, sex is det er mined by differ ences in t he chr omosomes.
In honeybees, eggs eit her ar e fer t ilized wit h a sper m and become diploid females, or
t hey ar e not fer t ilized and become haploid males, called drones.
• Female gr asshopper s have t wo X chr omosomes, and males have just one. The sper m
det er mines t he sex of t he zygot e. If a sper m wit hout an X fer t ilizes an egg, t he zygot e
becomes a male gr asshopper .
• Humans have differ ent sex chr omosomes, X and Y. Males have X and Y; females
have X and X.
• Sex of t he offspr ing is det er mined by t he sper m.
• Females do not have a Y chr omosome, so all nor mal female gamet es (eggs) have one
X chr omosome. Ma les ha ve one X chr omosome a nd one Y chr omosome, so ha lf of t he
male gamet es (sper m) have an X chr omosome and t he ot her half have a Y chr omosome.
If a sper m wit h an X chr omosome r eaches t he egg, t he r esult ing offspr ing will be
female (XX). If a sper m wit h a Y chr omosome r eaches t he egg, t he r esult ing offspr ing
will be male (XY).
• In humans, maleness is det er mined by t he pr esence of t he Y chr omosome.
The X and Y chromosomes have different functions
• Th e gen e t h a t det er mi n es ma l en es s wa s i den t i fi ed by s t u dyi n g peopl e wi t h
chr omosomal abnor malit ies.
• Some XY females wer e found. Some XX males wer e found. The XY females had a
piece missing fr om t he Y, and t he XX males had a piece of a Y on one X. The fr agment
missing fr om t he Y chr omosome in XY females or pr esent on t he X chr omosome in
XX males cont ained t he maleness-det er mining gene. The gene was named S RY (for
sex-det er mining region on t he Y chr omosome). The S RY gene codes for a funct ional
pr ot ein involved in pr imar y sex det er minat ion. If t his pr ot ein is pr esent , t est es develop;
if not , ovar ies develop. A gene on t he X chr omosome called DAX1 pr oduces an ant i-
t est is fact or . The S RY gene pr oduct in a male inhibit s t he gene DAX1, and no maleness
inhibit or is made.
• Secondar y sex t r ait s like br east development , body hair , and voice ar e influenced by
hor mones such as t est ost er one and est r ogen.
• Drosophila chr omosomes follow t he same pat t er n as humans, but t he mechanism is
differ ent . The males ar e XY, and females XX. Th e r a t i o of X ch r omos ome s t o t h e
a u t os oma l s e t s d e t e r mi n e s s e x . Two X chr omosomes for each diploid set yield
females. One X for each diploid set yields males. (XO is st er ile; XY is fer t ile).
• Bir ds, mot hs, and but t er flies have XX males and XY females. These ar e called ZZ
males and ZW females t o help pr event confusion. In t hese or ganisms, t he egg r at her
t han t he sper m det er mines t he sex of t he offspr ing.
Genes on sex chromosomes are inherited in special ways
• The Y car r ies ver y few genes (about 20 ar e known), but t he X car r ies a gr eat var iet y
of char act er s. Females wit h XX ar e diploid for X-linked genes; males wit h XY ar e
ha ploid. This pa r t ia l ha ploid condit ion of sex chr omosomes for ma les is ca lled
hemizygous. This gener at es a special t ype of inher it ance called sex-linked inher it ance.
Genetics 211
Human beings display many sex-linked characters
• The human X chr omosome car r ies t housands of genes. The probability for a male of
having an X-linked genetic disease caused by a mutant recessive allele is much higher
than it is for a female. Barring inbreeding, the probability of a woman having a recessive
X-linked genetic disease is the square of frequency of the disease-causing allele. (That
is the probability that she would inherit the gene from her mother times the probability
t hat she would inherit it from her fat her.) Because men have only one X chromosome,
and t hey express what t hey get whet her it is dominant or recessive, t he probabilit y for
a man of having an X-linked recessive genetic disease is simply the frequency of the
allele for the disease in the population. The number of let hal or severely det riment al
genes on t he X is kept low by t he hemizygous st at e of t he males.
• Pedigr ee analysis of X-linked r ecessive phenot ypes r eveal cer t ain pat t er ns: The
phenot ype appear s much mor e oft en in males t han in females. A male wit h t he
mut at ion can pass it on only t o his daught er s, t hr ough an X-bear ing sper m; his sons
get his Y chr omosome, which does not car r y t he t r ait . Daught er s who r eceive one
mut ant X ar e het er ozygous car r ier s. They pass t he allele t o appr oximat ely half of
t heir sons and daught er s. The mut ant phenot ype can skip a gener at ion if t he mut at ion
is passed fr om a male t o his daught er and t hen t o her son. The most common for ms
of mu s cu l a r d ys t r op h y a n d h e mop h i l i a , a s we l l a s r e d -gr e e n col or b l i n d n e s s ,
ar e a few X-linked human phenot ypes.
• Contrary to popular opinion, male pattern baldness is not X-linked. This is a sex-
i n f l u en ced t r a i t that is probably subject to hormonal influence. Males require only
one gene for baldness to appear; female baldness requires the presence of two genes.
Cytoplasmic Inheritance: Extra chromosomal Inheritance
• Mendelian genet ics is t he genet ics of t he nucleus, yet ot her cyt oplasmic or ganelles
car r y genet ic mat er ial. Mit ochondr ia, chlor oplast s, and ot her plast ids possess a small
amount of DNA. Humans have about 600,000 genes in t he nucleus and 37 genes in
mit ochondr ia.
• Plast id genomes ar e five t imes lar ger t han t hose of mit ochondr ia. The genome is t he
t ot al configur at ion of genet ic mat er ial. Mit ochondr ia and plast ids ar e passed on by
t he mot her only, a s t he egg cont a ins a bunda nt cyt opla sm a nd or ga nelles. The
mit ochondr ia in sper m do not t ake par t in gamet e union. Some chlor oplast s ar e
whit e, not gr een, because of a mut at ion in t heir DNA. Mit ochondr ial mut at ions may
also be linked t o human genet ic diseases.
• Mit ochodr ia is r esponsible for cyt oplasmic Male st er lit y in Maize
One Gene, One Polypeptide
• Ther e ar e many st eps bet ween genot ype and phenot ype; genes cannot by t hemselves
dir ect ly pr oduce a phenot ype. A gene is defined as a DNA sequence.
• In t he 1940s, Beadle and Tat um showed t hat an alt er ed gene r esult ed in an alt er ed
phenot ype t hat showed up as an alt er ed enzyme.
• They used t he br ead mold Neurospora crassa. This is an or ganism wit h a haploid
veget at ive life cycle, so r ecessive mut at ions ar e easy t o det ect . Neurospora wer e
gr own on minimal medium consist ing of just sucr ose, miner als, and a few vit amins.
212 CSIR-NET Life Sciences
Wild-t ype Neurospora wer e t r eat ed wit h a mut agen, an agent t hat causes changes in
t he DNA. Aft er t r eat ment , t hey wer e gr own in a complet e medium. When t est ing
some of t he t r eat ed st r ains, some wer e found t hat could no longer gr ow on minimal
medium, but inst ead needed cer t ain supplement s. These nut r ient -r equir ing auxot r ophs
wer e assumed t o have mut at ed. For each auxot r ophic st r ain, Beadle and Tat um wer e
able t o find a single compound t hat could suppor t it s gr owt h. One gr oup of mut ant s
needed ar ginine t o gr ow. Mapping st udies est ablished t hat some of t hese arg mut at ions
a r e a t differ ent loci, a nd t her efor e a r e in differ ent genes. Bea dle a nd Ta t um
demonst r a t ed t ha t t hese differ ent mut a nt s ha d defect ive genes for t he sa me
biochemical pat hway, t he pat hway leading t o ar ginine synt hesis.
• If t he gene defect affect ed ear lier enzyme st eps in t he pat hway, sever al differ ent
subst ances could subst it ut e for ar ginine. If t he defect was for t he enzyme st ep just
befor e ar ginine synt hesis, only ar ginine could subst it ut e. Beadle and Tat um t hus
post ulat ed t he one gene, one enzyme hypot hesis .
• Lat er it was lear ned t hat some enzymes ar e composed of differ ent subunit s coded for
by separ at e genes. The one gene, one enzyme hypot hesis was lat er changed t o t he one-
gene, one-polypept ide hypot hesis. Even t his hypot hesis r equir es modificat ion because
some genes code for RNA molecules t hat ar e never t r anslat ed int o polypept ides.
MUTATIONS: Heritable Changes in Genes
• Mut at ions ar e her it able changes in DNA—changes t hat ar e passed on t o daught er
cells. In single-celled or ganisms, any mut at ions t hat occur ar e passed t o t he daught er
cells at t he t ime of cell division.
• Mult icellular or ganisms have t wo t ypes of mut at ions:
• Somat ic mut at ions ar e passed on dur ing mit osis, but t he affect ed cells never become
gamet es and so do not pass t o subsequent gener at ions and
• Ger m-line mut at ions ar e mut at ions t hat occur in cells t hat might give r ise t o gamet es.
• Some mut at ions cause visible phenot ypic change. Ot her s cause met abolic changes
t hat might not yet be det ect able.
• Some mut at ions exer t t heir effect only under cer t ain r est r ict ive condit ions. These
ar e called con d i t i on a l mu t a n t s . They ar e unaffect ed under per missive condit ions,
but expr ess t he mut ant phenot ype at t he r est r ict ive condit ion. Temper at ur e-sensit ive
mut ant s ar e an example.
• All mut at ions ar e alt er at ions of t he DNA nucleot ide sequence and ar e of t wo t ypes:
Point mut at ions ar e mut at ions of single genes and Chr omosomal mut at ions ar e
changes in t he ar r angement s of chr omosomal DNA segment s.
Point mutations are changes in single bases
• Point mut at ions r esult fr om t he addit ion or subt r act ion of a nucleot ide base or t he
subst it ut ion of one base for anot her . Point mut at ions can occur as a r esult of mist akes
dur ing DNA r eplicat ion, or by envir onment al mut agens, such as chemicals and
r adiat ion. Because of r edundancy in t he genet ic code, some point mut at ions r esult in
no change in t he amino acids in t he pr ot ein. Th e s e a r e ca l l e d s i l e n t mu t a t i on s .
• Some mut a t ions ca use a n a mino a cid subst it ut ion. These a r e ca lled mi s s e n s e
mu t a t i on s . An example in humans is sickle-cell anemia, a defect in t he b-globin
Genetics 213
subunit s of hemoglobin. The r ed blood cells collapse when oxygen levels ar e low.
Missense mut at ions might r educe t he funct ioning of a pr ot ein or disable it complet ely.
• Non s e n s e mu t a t i on s ar e base subst it ut ions t hat cause a change fr om a codon t hat
inst r uct s t he incor por at ion of an amino acid t o a codon t hat t er minat es t r anslat ion.
• A fr a me -s h i ft mu t a t i on is when a single base is inser t ed or delet ed in a gene. This
causes t he most disr upt ion when t he event occur s at or near t he beginning of t he
t emplat e. This t ype of mut at ion shift s t he code, changing many of t he codons t o
differ ent codons. These shift s almost always lead t o t he pr oduct ion of nonfunct ional
pr ot eins.
Chromosomal mutations are extensive changes in the genetic material
• DNA molecules can br eak and r e-for m. This can cause four differ ent t ypes of mut at ions:
delet ions, duplicat ions, inver sions, and t r anslocat ions. Delet ions ar e a loss of a
chr omosomal segment . Duplicat ions ar e a r epeat of a segment . Br eaking and r ejoining
l ea ds t o i n ver s i on s i f s egmen t s get r ea t t a ch ed i n t h e oppos i t e or i en t a t i on .
Tr a nsloca t ions r esult when a por t ion of one chr omosome a t t a ches t o a not her .
Tr anslocat ions can be r ecipr ocal or nonr ecipr ocal. Tr anslocat ions can make synapses
in meiosis difficult and can lead t o aneuploidy (t oo many or t oo few chr omosomes).
Human Disorders due to Chromosome Alterations
• Tr i s omy 21 (Down Syn d r ome ): Affect s on aver age one out of 700 childr en bor n in
USA. individuals ar e shor t in st at ur e, suffer var ying degr ees of ment al r et ar dat ion
and ar e suscept ible t o a var iet y of diseases (leukemia, Alzheimer ’s) Higher incidence
of Down’s childr en bor n t o mot her s over 35 year s old.
• Tr i s omy 18 (Ed wa r d Syn d r ome )
• Tr i s omy 13 (P a t a u Syn d r ome )
• Kli n e fe lt e r Syn d r ome (XXY): occur s one in ever y 2000 bir t hs. male sex or gans,
but individuals ar e st er ile and have feminine body char act er ist ics
• Tu r n e r Syn d r ome (XO): fema les a r e mon os omic for X ch r omos ome, n or ma l
int elligence but do not fully develop secondar y sex char act er ist ics ar e puber t y., shor t
in st at ur e
Structural Alterations
• “Cr i d u Ch a t ” is r esult of delet ion of a r egion of chr omosome 5: Individuals ar e
ment ally r et ar ded and have a cr y t hat sounds like a cat .
• Chr omosomal Tr anslocat ions: associat ed wit h some diseases e.g.. Tr anslocat ion
of a par t of chr omosome 22 wit h a par t of chr omosome 9 r esult s in chr onic
myelogenous leukemia (P h i l a d e l p h i a 22).
Mutations can be spontaneous or induced
• Spont aneous mut at ions ar e per manent changes t hat occur wit hout out side influence.
Spont aneous mut at ions may be caused by any of sever al mechanisms. Nucleot ides
occa siona lly cha nge t heir st r uct ur e (ca lled a t a u t ome r i c s h i ft ). A ba se ma y
t empor ar ily change t o it s unusual t aut omer at t he same t ime t hat r eplicat ion is
occur r ing. The t aut omer may pair wit h t he alt er nat e pur ine if it is a pur ine, or t he
alt er nat e pyr imidine if it is a pyr imidine.
214 CSIR-NET Life Sciences
• DNA polymer ase somet imes makes er r or s in r eplicat ion. These er r or s ar e oft en
r epair ed by t he pr oofr eading funct ion of t he r eplicat ion complex, but some er r or s
escape and become per manent .
• Meiosis is imper fect . Nondisjunct ion can occur . Random chr omosome br eaks
r ejoin incor r ect ly, leading t o t r anslocat ions.
• Induced mut at ions ar e per manent changes caused by some out side agent .
• Some chemicals alt er covalent bonds in nucleot ides. Nit r ous acid deaminat es
cyt osine, conver t ing it t o ur acil. DNA polymer ase mist akes ur acil for t hymine
and put s an A in dur ing r eplicat ion inst ead of t he G t hat would have been
incor por at ed ot her wise.
• Benzoapyr ene, a pr oduct of incomplet e combust ion, which is found in all smoke,
adds a lar ge chemical gr oup t o guanine, making it unavailable for base pair ing.
Any base might be inser t ed t o fill t he gap.
• Radiat ion damages DNA. Ionizing r adiat ion (X r ays) pr oduces highly r eact ive
compounds and at oms called fr ee r adicals. Gamma rays also produce free radicals.
Fr ee r adicals can alt er bases or br eak t he sugar –phosphat e backbone, causing
chr omosomal abnor malit ies. Ult r aviolet r adiat ion is absor bed by pyr imidines in
t he DNA, and when t wo t hymines or t wo cyt osines ar e next t o each ot her on t he
same st r and of a double-st r anded DNA molecule, a covalent bond can for m.
Their int er st r and covalent bonds make t he DNA unr eplicable.
• The long-t er m benefit of mut at ions is t hat t hey pr ovide a genet ic diver sit y for
evolut ion and account for all t he differ ences bet ween and wit hin or ganisms,
excluding t he effect of differ ent envir onment s. The det r iment of mut at ion is t he
out r ight deat h or poor fit of an or ganism t o it s envir onment .
Mutations are the raw material of evolution
• Mut at ions ar e r ar e event s and most of t hem ar e point mut at ions involving one
nucleot ide. Fr equency of mut at ions is much lower t han one mut at ion per 10
4
genes
per DNA duplicat ion. Somet imes t hey ar e as r ar e as one per 10
9
genes per duplicat ion.
Differ ent or ganisms var y in mut at ion fr equency.
• Mut at ions can be det r iment al, neut r al, or occasionally beneficial. Humans have 1,000
t imes t he DNA of a pr okar yot e. This is at least par t ially due t o duplicat ion of DNA
sequences, and t hen t o diver gence of t he sequences over t ime. Random accumulat ion
of mut at ions in t he ext r a copies of genes can lead t o t he pr oduct ion of new useful
pr ot eins.
Mutation and Variation
• Genet ic var iat ion r efer s t o differ ences in genot ype among or ganisms. Genot ypes can
be classified accor ding t o inher it ance pat t er ns
• n u clea r gen es : inher it ed t hr ough bot h male or female gamet e
• ma t e r n a l l y i n h e r i t e d : mit ochondr ia in animals and chlor oplast in angiosper ms.
• p a t e r n a lly i n h e r i t e d : chlor oplast s in conifer s.
• Envir onment al var iat ion: differ ences in t he envir onment due t o phenot ypic plast icit y.
Genetics 215
• Mat er nal effect s: char act er ist ics of a mot her s offspr ing due t o nongenet ic-influences,
e.g., via deposit ion of yolk in eggs. Differ ences among offspr ing ar e a r eflect ion of
genot ypic/envir onment al var iat ion in t he mot her .
Since only genet ic var iat ion is her it able, one needs t o have means t o dist inguish bet ween
genet ic and ot her sour ces of var iat ion.
• For t rait s influenced by few genes cr osses t o pr oduce F1, F2, and backcr osses may
yield Mendelian r at ios, and t hus indicat e genet ic var iat ion.
• For polygenic t rait s, cor r elat ions bet ween (i) par ent s and offpr ing, (ii) among siblings,
indicat e genet ic var iat ion. To exclude t he possibilit y of mat er nal effect s, one can,
e.g., det er mine t he cor r elat ion bet ween fat her s and t he offspr ing. To exclude t he
possibilit y t hat t he obser ved cor r elat ions ar e due t o a shar ed envir onment , one can
eit her (i) r andomize t he offspr ings’ envir onment (e.g., dist r ibut e bir d’s eggs among
differ ent fost er par ent s) or (ii) r ear t he or ganisms for sever al gener at ions in t he
sa me envir onment , a nd obser ve pa r ent offspr ing cor r ela t ions (common-garden
experiment s).
Sources of genetic variation
Her e, sour ces of var iat ion less likely t o be discussed in ot her cour ses ar e emphasized.
• P oi n t mu t a t i on s Tr ansit ions (C <-> T, A <->G) Tr ansver sions ( A <-> C, A <-> T,
G<-> C, G<-> T)
In coding r egions point mut at ions can be synonymous or non-synonymous, be missense
or nonsense mut at ions. Mut at ion r at es ar e of t he or der of 10
-9
per base pair per
gener at ion.
Insertions/Deletions
May lead t o fr ame shift mut at ions and gene duplicat ions.
• r eplicat ion slippage: shor t indels
• unequal cr ossing over : longer indels
Recombination
homologous r ecombinat ion in meiosis
• r ecipr ocal r ecombinat ion
• gene conver sion (in fungi)
• non-homologous r ecombinat ion (unequal cr ossing over )
Chromosome rearrangements
polyploidizat ion
• duplicat ions/delet ions
• inver sions -per icent r ic (including t he cent r omer e) & par acent r ic
• fi s s i on s /fu s i on s (a r e a s p e ci a l ca s e of t r a n s l oca s t i on s i n vol vi n g on e
minichr omosome
• (r ecipr ocal) t r anslocat ions
All t hese ar e impor t ant because of posit ion effect s. Pr ominent ca uses of chr omosome
r ear r angement s ar e unequal cr oss-over and t r ansposable element act ivit y (see below). The
r at e of any given class of r ear r angement s is high, 10
–3
– 10
–4
per gamet e per gener at ion.
216 CSIR-NET Life Sciences
TRANSPOSITION
Th r ee ma i n ki n ds of t r a n s pos i t i on a r e n on -r epl i ca t i ve t r a n s pos i t i on , r epl i ca t i ve
t r ansposit ion, and r et r oposit ion. In all t hr ee, a duplicat ion (dir ect r epeat ) in donor DNA is
gener at ed t hat is a few base pair s long.
Classes of transposable elements
• I n s e r t i on s e q u e n ce s (Se l fi s h DNA): (ISs) ar e t he simplest t r ansposable element s
and ar e abundant in bact er ia, bact er iophages, plasmids, maize (t he mobile element s
char act er ized by B. McClint ock). They car r y only t he t r ansposase funct ion, e.g., IS1 :
insA and insB encode for t r ansposase, flanked by inver t ed r epeat s.
• Tr a n s p os on s (J u mp i n g ge n e s ): similar t o IS, but t hey car r y addit ional genes which
fr equent ly confer : a nt ibiot ic r esist a nce, hea vy met a l r esist a nce, hea t r esist a nce
Examples, cat is chlor amphenicol r esist ance, bla is st r ept omycin r esist ance). Some
t r ansposons, such as Tn 9 ar e compound t r ansposons consist ing of mor e t han one IS
sequ en ce (Her e, t h e en t ir e elemen t ca n t r a n spose or on ly t h e I S sequ en ces).
Tr ansposons ar e ver y widespr ead, e.g., Drosophila cont ains 50 – 100 kinds. The
bact er iophage Mu is a t r ansposon t hat also cont ains genes for phage packaging.
• The pr ot ot ype Dr osophila t r ansposons ar e called cop ia , because it is pr esent in copious
quant it y. Similar t r ansposable element s in yeast ar e called Ty for ‘Tr ansposon yeast ’,
ot her t ype pet it e also occur . They ar e named as AC-Ds (Act ivat or -Dissociat or ) element s
in maize.
• The evolut ionar y r elevance of t r ansposons is under scor ed by t he phenomenon of
hybrid dysgenesis a ssocia t ed wit h P-element s in Drosophila. P -e l e me n t s i n D.
melanogast er ar e <40 yr s old, and have pr obably been t r ansfer r ed via hor izont al
t r ansfer fr om anot her species. In hybr id dysgenesis, P element s become act ivat ed
only in t he ger mline, and only in par t icular t ypes of cr osses. They gener at e many
mut at ions t hat cause failur e of t he ger mline t o develop. In somat ic cells, a r epr essor
is pr oduced by alt er nat ive splicing t hat eliminat es t he mobilit y of P-element s. P-
element s may pr ovide mechanisms for r epr oduct ive isolat ion in fr uit flies.
Retroelements
All element s t hat car r y r ever se t r anscr ipt ase, whet her t r ansposable or not . They fall int o
sever al cat egor ies.
• R et r ovi r u ses
Their minimal gene cont ent consist s of t he gag gene encoding t he vir al capsid, t he
pol gene encoding r eplicat ion funct ions including r ever se t r anscr ipt ase, and t he env
gene encoding t he envelope.
• Retrotransposons
They do not have an env gene, t hus t hey cannot under go pr oper packaging. Examples
ar e copia, gypsy, and Ty1
• Retroposons
Like r et r ot r ansposons, but t hey lack LTRs. Examples include LINEs (long int er sper sed
nucleot ide element s) in mammals.
Genetics 217
• Retrons
They have no LTRs, and encode only t he r ever se t r anscr ipt ion funct ion. They occur
in bact er ia and in t he mit ochondr ial genome of plant s, do not excise, and t hus for m
par t of t he genome.
• (Processed) retrosequences
Ar e genomic DNA sequences t hat ar e similar t o ot her , int r on-car r ying genes, but ar e
char act er ized by lack of int r ons, boundar ies cor r esponding t o t r anscr ibed r egions of a
gene, st r et ches of poly-A at 3'-end , shor t dir ect r epeat s at bot h ends, indicat ing past
t r anspot sit ion and no linkage wit h t he or iginal gene. The pr ocess of gener at ing
pr ocessed r et r osequences involves r ever se t r anscr ipt ase.
• They ar e oft en nonfunct ional, because r ever se t r anscr ipt ase has a high er r or r at e, no
r egulat or y signals ar e t r ansposed & t heir 5' end is t r uncat d. Such non-funct ional
r et r osequ en ces a r e ca lled r e t r op s e u d oge n e s . Exa mples include SI NE (shor t
int er sper sed nucleot ide element s), LINE, and Alu element s (Alu is a r et r osequence
der ived fr om t he 7SL RNA gene which is a par t of t he signal r ecognit ion par t icle.
• Some of t hese element s have acquir ed addit ional sequences, e.g., LINE element s
may have a r ever se t r anscr ipt ase. Funct ional r et r osequences also exist . They ar e
called r et r ogen es . Examples ar e human phosphoglycer at e kinase and muscle-specific
calmodulin gene in chicken.
Evolutionary effects of transposable elements
They ccnfer var ious r esist ances, cause an incr ease in host genome size, cause alt er at ion of
host gene expr ession, eit her by inser t ing int o cont r ol r egions, or int o coding r egions. Example:
The humen t het a1 globin gene has a t r uncat ed Alu sequence as par t of it s pr omot er . Elevat e
t he mut at ion r at e of host genes, cause chr omosome r ear r angement s: delet ions/duplicat ions,
inver sions, and t r anslocat ions. These ar e somet imes mediat ed by t r ansposit ion it self (t wo
t r ansposable element s wit h int er vening DNA t r ansposing as a unit ) & unequal cr oss-over
bet ween r epeat ed copies of t r ansposable element s, which may duplicat e genes in t he r egion.
Unequal cr oss-over may cause changes in t he domain st r uct ur e of genes via unequal cr oss-
over . For example, a mut at ion of t he low-densit y-lipopr ot ein gene in humans causing high
cholest er ol levels is t hought t o have or iginat ed by unequal cr ossing-over involving Alu sequences.
DNA: The Genetic Material —The physical carrier of inheritance
• Fr i ed r i ch Mei sch er in 1869 isolat ed DNA fr om fish sper m and t he pus of open
wounds. Since it came fr om nuclei, Meischer named t his new chemical, nuclein.
Subsequent ly t he name was changed t o nucleic acid and last ly t o deoxyr ibonucleic
acid (DNA). Rober t Feulgen, in 1914, discover ed t hat fuchsin dye st ained DNA. DNA
was t hen found in t he nucleus of all eukar yot ic cells.
• Dur ing t he 1920s, biochemist P .A. Le ve n e analyzed t he component s of t he DNA
molecule. He found it cont ained four nit r ogenous bases: cyt osine, t hymine, adenine,
and guanine; deoxyr ibose sugar ; and a phosphat e gr oup. He concluded t hat t he basic
unit (nucleot ide) was composed of a base at t ached t o a sugar and t hat t he phosphat e
also at t ached t o t he sugar . He (unfor t unat ely) also er r oneously concluded t hat t he
pr opor t ions of bases wer e equal and t hat t her e was a t et r anucleot ide t hat was t he
r epea t ing st r uct ur e of t he molecule. The nucleot ide, however , r ema ins a s t he
218 CSIR-NET Life Sciences
fundament al unit (monomer ) of t he nucleic acid polymer . Ther e ar e four nucleot ides:
t hose wit h cyt osine (C), t hose wit h guanine (G), t hose wit h adenine (A), and t hose
wit h t hymine (T).
• Molecular st r uct ur e of t hr ee nir ogenous bases. In t his diagr am, t her e ar e t hr ee
phosphat es inst ead of t he single phosphat e found in t he nor mal nucleot ide.
DNA as Genetic Material
• Dur ing t he ear ly 1900s, t he st udy of genet ics began in ear nest : t he link bet ween
Mendel’s wor k and t hat of cell biologist s r esult ed in t he chr omosomal t heor y of
i n h er i t a n ce; Ga r r o d pr oposed t h e lin k bet ween gen es a n d “in bor n er r or s of
met abolism”; and t he quest ion was for med: what is a gene? The answer came fr om
t he st udy of a deadly infect ious disease: pneumonia.
• Dur ing t he 1920s Fr ed er i ck Gr i ffi t h st udied t he differ ence bet ween a disease-causing
st r ain of t he pneumonia causing bact er ia (S t rept ococcus peumoniae) and a st r ain t hat
did not cause pneumonia. The pneumonia-causing st r ain (t he S st r ain) was sur r ounded
by a capsule. The ot her st r ain (t he R st r ain) did not have a capsule and also did not
cause pneumonia. Fr eder ick Gr iffit h (1928) was able t o induce a nonpat hogenic st r ain
of t he bact er ium S t rept ococcus pneumoniae t o become pat hogenic. Gr iffit h r efer r ed
t o a t r ansfor ming fact or t hat caused t he non-pat hogenic bact er ia t o become pat hogenic.
Gr iffit h inject ed t he differ ent st r ains of bact er ia int o mice. The S st r ain killed t he
mice; t he R st r ain did not . He fur t her not ed t hat if heat killed S st r ain was inject ed
int o a mouse, it did not cause pneumonia. When he combined heat -killed S wit h Live
R and inject ed t he mixt ur e int o a mouse (r emember neit her alone will kill t he mouse)
Genetics 219
t hat t he mouse developed pneumonia and died. Bact er ia r ecover ed fr om t he mouse
had a capsule and killed ot her mice when inject ed int o t hem.
Hypotheses
1. The dead S st r ain had been r eanimat ed/r esur r ect ed.
2. The Live R had been t r ansfor med int o Live S by some “t r ansfor ming fact or ”.
Fur t her exper iment s led Gr iffit h t o conclude t hat number 2 was cor r ect .
• I n 1944, Os wa ld Ave r y, Coli n Ma cLe od , a n d Ma clyn McCa r t y r evisit ed Gr iffit h’s
exper iment and concluded t he t r ansfor ming fact or was DNA. Their evidence was
st r ong but not t ot ally conclusive. The t hen-cur r ent favor it e for t he her edit ar y mat er ial
was pr ot ein; DNA was not consider ed by many scient ist s t o be a st r ong candidat e.
• The br eakt hr ough in t he quest t o det er mine t he her edit ar y mat er ial came fr om t he
wor k of Ma x De l b r u ck a n d Sa l va d or Lu r i a in t he 1940s. Bact er iophage ar e a t ype
of vir us t hat at t acks bact er ia, t he vir uses t hat Delbr uck and Lur ia wor ked wit h wer e
t h os e a t t a ck i n g Esch eri ch i a col i , a ba ct er i u m fou n d i n h u ma n i n t es t i n es .
Bact er iophages consist of pr ot ein coat s cover ing DNA. Bact er iophages infect a cell by
inject ing DNA int o t he host cell. This vir al DNA t hen “disappear s” while t aking over
t he bact er ial machiner y and beginning t o make new vir us inst ead of new bact er ia.
Aft er 25 minut es t he host cell bur st s, r eleasing hundr eds of new bact er iophage. Phages
have DNA and pr ot ein, making t hem ideal t o r esolve t he nat ur e of t he her edit ar y
mat er ial.
• In 1952, Al fr e d D. He r s h e y a n d Ma r t h a Ch a s e conduct ed a ser ies of exper iment s
t o det er mine whet her pr ot ein or DNA was t he her edit ar y mat er ial. By labeling t he
DNA and pr ot ein wit h differ ent (and mut ually exclusive) r adioisot opes, t hey would be
able t o det er mine which chemical (DNA or pr ot ein) was get t ing int o t he bact er ia.
Such mat er ial must be t he her edit ar y mat er ial (Gr iffit h’s t r ansfor ming agent ). Since
DNA cont ains Phosphor ous (P) but no Sulfur (S), t hey t agged t he DNA wit h r adioact ive
Phosphor ous-32. Conver sely, pr ot ein lacks P but does have S, t hus it could be t agged
wit h r adioact ive Sulfur -35. Her shey and Chase found t hat t he r adioact ive S r emained
out side t he cell while t he r adioact ive P was found inside t he cell, indicat ing t hat DNA
was t he physical car r ier of her edit y.
Diagr ams illlust r at ing t he Her shey and Chase exper iment t hat suppor t ed DNA as
t he her edit ar y mat er ial while it also showed pr ot ein was not t he her edit ar y mat er ial.

220 CSIR-NET Life Sciences
GENETIC DISORDERS
Human Allelic Disorders (Recessive)
• The fir st Mendelian t r ait in humans was descr ibed in 1905 (br achydact ly) by Dr .
Far abee (no r elat ion t o your aut hor ). Now mor e t han 3500 human genet ic t r ait s ar e
known.
• Al b i n i s m, t he lack of pigment at ion in skin, hair , and eyes, is also a Mendelian
human t r ait . Homozygous r ecessive (aa) individuals make no pigment s, and so have
face, hair , and eyes t hat ar e whit e t o yellow. For het er ozygous par ent s wit h nor mal
pigment at ion (Aa), t wo differ ent t ypes of gamet es may be pr oduced: A or a. Fr om
such a cr oss 1/4 of t he childr en could be albinos. The br own pigment melanin cannot
be made by albinos. Sever al mut at ions may cause albinism: 1) t he lack of one or
anot her enzyme along t he melanin-pr oducing pat hway; or 2) t he inabilit y of t he enzyme
t o ent er t he pigment cells and conver t t he amino acid t yr osine int o melanin.
• P h e n yl k e t on u r i a (P KU) is r ecessively inher it ed disor der whose suffer er s lack t he
abilit y t o synt hesize an enzyme t o conver t t he amino acid phenylalanine int o t yr osine.
Individuals homozygous r ecessive for t his allele have a buildup of phenylalanine and
abnor mal br eakdown pr oduct s in t he ur ine and blood. The br eakdown pr oduct s can
be har mful t o developing ner vous syst ems and lead t o ment al r et ar dat ion. 1 in 15,000
infant s suffer s fr om t his pr oblem. PKU homozygot es ar e now r out inely t est ed for in
most st a t es. I f you look closely a t a pr oduct cont a ining Nut r a -sweet a r t ificia l
sweet ener , you will see a war ning t o PKU suffer er s since phenylalanine is one of t he
amino acids in t he sweet ener . PKU suffer er s ar e placed on a diet low in phenylalanine,
en ou gh for met a bolic n eeds bu t n ot en ou gh t o ca u se t h e bu ildu p of h a r mfu l
int er mediat es.
• Ta y-Sa ch s Di sea se is an aut osomal r ecessive r esult ing in degener at ion of t he ner vous
syst em. Sympt oms manifest aft er bir t h. Childr en homozygous r ecessive for t his allele
r ar ely sur vive past five year s of age. Suffer er s lack t he abilit y t o make t he enzyme N-
acet yl-hexosaminidase, which br eaks down t he GM2 ganglioside lipid. This lipid
accumulat es in lysosomes in br ain cells, event ually killing t he br ain cells. Alt hough
Genetics 221
r ar e in t he gener al populat ion (1 in 300,000 bir t hs), it was (unt il r ecent ly) higher (1 in
3600 bir t hs) among J ews of east er n cent r al Eur opean descent . One in 28 Amer ican
J ews is t hought t o be a car r ier , since 90% of t he Amer ican J ewish populat ion emigr at ed
fr om t hose ar eas in Eur ope. Most Tay-Sachs babies bor n in t he US ar e bor n t o non-
J ewish pa r ent s, who did not under go t est ing pr ogr a ms t ha t most US J ewish
pr ospect ive par ent s had.
• Si ck le -ce ll a n e mi a is an aut osomal r ecessive we have discussed in ot her sect ions.
Nine-per cent of US blacks ar e het er ozygous, while 0.2% ar e homozygous r ecessive.
The r ecessive allele causes a single amino acid subst it ut ion in t he bet a chains of
hemoglobin. When oxygen concent r at ion is low, sickling of cells occur s. Het er ozygot es
make enough “good bet a-chain hemoglobin” t hat t hey do not suffer as long as oxygen
concent r at ions r emain high, such as at sea-level.
• Cys t i c fi b r os i s is common in whit es of Eur opean descent (1 in 2500 affect ed), one
out of 25 whit es is a car r ier (4%), gene codes for chlor ide channel (defect ive gene due
t o addit ion or deler t ion of ext r a TTT sequence r esult s in mucous buildup in lungs)
• Pat ient s wit h Al ca p t on u r i a excr et ed copious amount of homogenit isic acid, which
has st ar t ling effect of color ing t heir ur ine black.
Human Allelic Disorders (Dominant)
• Aut osomal dominant s ar e r ar e, alt hough t hey ar e (by definit ion) mor e commonly
expr essed.
• Ac h on d r op l a s t i c d wa r fi s m occur s, even t hough suffer er s have r educed fer t ilit y.
• Hu n t i n gt on ’s d i s e a s e (also r efer r ed t o as Woody Gut hr ie’s disease, aft er t he folk
singer who died in t he 1960s) is an aut osomal dominant r esult ing in pr ogr essive
dest r uct ion of br ain cells. If a par ent has t he disease, 50% of t he childr en will have it
(unless t hat par ent was homozygous dominant , in which case all childr en would have
t he disease). The disease usually does not manifest unt il aft er age 30, alt hough some
inst ances of ear ly onset phenomenon ar e r epor t ed among individuals in t heir t went ies.
-Incur able; br ain det er ior at ion leads t o deat h. Gene is pr esent on ch r omos ome n o 4
and t her e is a d d i t i on of ext r a CAG sequ en ces (n or ma lly 25 r ep ea t s a r e p r esen t )
at end of h u n t i gt i n gene.
• Ne u r ofi b r oma t os i s : It is one of t he most common aut osomal dominant genet ic
diseases in human. It is of t wo t ypes namely NF1 and NF2. Individuals wit h t his
disease develop sm all-pigmented skin lesions called ca f é-a u -l a i t spot s and small soft
fleshy gr owt h called n e u r ofi b r oma t a , wh i ch ar e benign t umor s. These appear dur ing
adolescence and t end t o incr ease wit h t he age. The defect ive gene is NF1 locat ed on
chr omosome 17 abd codes a pr ot ein n e u r ofi b r omi n which act s as a t umor suppr essor
pr ot ein while NF2 gene is locat ed on chr omosome 22 and encodes a pr ot ein called
me r l i n which like NF1 gene pr oduct act as t umor suppr essor gene.
• P ol yd a c t l y is t he pr esence of a sixt h digit . In moder n t imes t he ext r a finger has
been cut off at bir t h and individuals do not know t hey car r y t his t r ait . One of t he
wives of Henr y VIII had an ext r a finger . In cer t ain sout her n families t he t r ait is also
mor e common. The ext r a digit is r ar ely funct ional and definit ely causes pr oblems
buying gloves, let alone fit t ing t hem on dur ing a mur der t r ial.
Sex-linked Traits
• Color b li n d n e s s afflict s 8% of males and 0.04 % of human females. Color per cept ion
depends on t hr ee genes, each pr oducing chemicals sensit ive t o differ ent par t s of t he
222 CSIR-NET Life Sciences
visible light spect r um. Red and gr een det ect ing genes ar e on t he X-chr omosome,
while t he blue det ect ion is on an aut osome.
• He mop h i l i a is a gr oup of diseases in which blood does not clot nor mally. Fact or s in
blood ar e involved in clot t ing. Hemophiliacs lacking t he nor mal Fact or VIII ar e said
t o have Hemophilia A, t he most common for m. Nor mal Fact or VIII can be supplied at
a high dollar and healt h r isk cost , alt hough t he development of biot echnologically
engineer ed Fact or VIII pr oduced by bact er ia lessens t he healt h r isk. England’s Queen
Vict or ia was a car r ier for t his disease. The allele was passed t o t wo of her daught er s
and one son. Since r oyal families in Eur ope commonly int er mar r ied, t he allele spr ead,
and may have cont r ibut ed t o t he downfall of t he Russian monar chy (Czar Nicholas’
son Alexei suffer ed fr om hemophilia A inher it ed fr om his mot her who car r ied Vict or ia’s
genet ic secr et ).
• Mu s cu l a r d ys t r op h y is a t er m encompassing a var iet y of muscle wast ing diseases.
The most common t ype, Du ch e n n e Mu s cu l a r Dys t r op h y (DMD), affect s car diac
and skelet al muscle, as well as some ment al funct ions. DMD is an X-linked r ecessive
occur r ing in 1 in 3500 newbor ns. Most suffer er s die befor e t heir 20t h bir t hday. In
1987, Louis Kunkel claimed t o have isolat ed a pr ot ein, dyst r ophin, pr esent in nor mal
individuals (about 0.002 % of t heir muscle pr ot ein) but absent in t wo individuals wit h
DMD. The lack of dyst r ophin is accompanied wit h a condit ion of muscle har dening
known as fibr osis, which r est r ict s blood supply t o t he muscle which t hen die.
• Ot h e r d i s e a s e s X-Li n k e d d i s e a s e include Le s ch -Nya n n s yn d r ome (Mut at ed
HGPTR gene) and F r a gi le X syn d r ome (Due t o fr agile sit e on X chr omosome at sit e
Xq27)
Se x l i mi t e d t r a i t s : The genes of t hese t r ait s ar e aut osomal and found in bot h sexes but
expr ess in one sex only e.g., milk glands in female, bear d in man, deep male voice, ant ler s in
male deer s, br illiant plumage in peacock, female or male musculat ur e et c. The expr ession of
t hese genes is affect ed by sex hor mones.
Se x i n fl u e n ce d t r a i t s : In cont r ast t o sex limit ed genes wher e t he expr ession of a t r ait is
limit ed t o one sex only, sex influenced genes ar e t hose aut osomal genes which ar e influenced
by t he sex of a bear er e.g., pat t er n of baldness, shor t index finger in male. These t r ait s appear
mor e fr equent ly in one sex t han in t he ot her . The baldness is affect ed by male hor mone
(t est ost er one).
• Some diseases ar e mu l t i fa c t or i a l which means t hey ar e due t o bot h genet ic and
envir onment al component s. Exa mp l e s : hear t disease, diabet es, cancer .
III. Technology for Genetic Testing
1. If t wo par ent s ar e car r ier s (het er ozygous) for a disease t hen t her e is a 1/4 chance t hat
t heir child will be affect ed (homozygous r ecessive).
• Car r ier s can be ident ified by molecular genet ic t est s.
• Fet al t est ing can be per for med by a mn i oce n t e s i s (14 t o 16 weeks gest at ion) or
ch or i on i c vi l l u s s a mp l i n g (10 weeks gest at ion).
• kar yot yping, biochemical, or molecular t est s can be per for med.
• Newbor n scr eening: Phenylket onur ia (PKU) t est is r out inely per for med on
newbor ns (affect s 1 out of 10,000 bir t hs). Affect ed individuals cannot br eakdown
phenylalanine and buildup t oxic levels of it s bypr oduct phenylpyr uvat e.
PRACTICE TEST PAPER-I
1. Which of t he following chr omosomal alt er at ions would you expect t o have t he most
dr ast ic consequences?
(a) inver sion (b) duplicat ion
(c) t r anslocat ion (d) delet ion
2. The most common let hal genet ic disease in t he Unit ed St at es is
(a) sickle-cell an emia (b) cyst ic fibr osis
(c) Hunt ingt on’s disease (d) hemophilia.
3. Ther e ar e var ious pr ocedur es t hat can be used t o det ect genet ic disor der s befor e
bir t h. Among t he t est s discussed in t his chapt er , ____ is t he least invasive, while
____ car r ies t he highest r isk.
(a) chor ionic villi sampling . . . amniocent esis
(b) ult r asound imaging . . . fet oscopy
(c) fet oscopy . . . chor ionic villi sampling
(d) fet oscopy . . . amniocent esis
4. Tay-Sachs disease r uns in Seema’s family. On a family pedigr ee, she saw a half-
dar kened cir cle. This r epr esent ed
(a) a male wit h Tay-Sachs (b) a female wit h Tay-Sachs
(c) a car r ier male (d) a car r ier female.
5. On a pedigr ee t r acing t he inher it ance of PKU, a hor izont al line joins a black squar e
and a half-black cir cle. What fr act ion of t his couple’s childr en would you expect t o
suffer fr om PKU?
(a) none (b) 1/4
(c) 1/2 (d) 3/4
6. Ram and Nidhi ar e appar ent ly nor mal, but t heir daught er was bor n wit h alkapt onur ia,
an inher it ed met abolic disor der . If alkapt onur ia is like most human her edit ar y
disor der s, t he pr obabilit y of t heir next child being bor n wit h alkapt onur ia is
(a) 0 (b) 1/4
(c) 1/2 (d) 2/3
7. Sever a l i n h er i t ed di s or der s a r e mu ch mor e common i n cl os e-kn i t r el i gi ou s
communit ies, such as t he Amish(J ews), t han in t he gener al populat ion. This is at
least par t ly due t o t he fact t hat
(a) people in such communit ies ar e mor e likely t o mar r y r elat ives
(b) shar ed envir onment al condit ions such as diet can incr ease mut at ion r at e
(c) moder n medical car e is not widely available in such communit ies
(d) communit y member s car e for each ot her and disor der s ar e possed on.
8. A her it able feat ur e is a ______ and may have t wo or mor e var iant s called______ .
(a) t r ait /char act er ist ics (b) char act er /t r ait s
(c) char act er /fact or s (d) t r ait /fact or s
224 CSIR-NET Life Sciences
9. In a cr oss bet ween t wo het er ozygot es (Aa), t he F
2
gener at ion will be
(a) in t he r at io 1:3 het er ozygous t o homozygous
(b) all het er ozygous
(c) in t he r at io 1:1 homozygous t o het er ozygous
(d) in t he r at io 1:3 homozygous t o het er ozygous
10. You set up an exper iment in which you br eed t wo populat ions of t r ue-br eeding pea
plant s. The fir st t r ue-br eeding populat ion has yellow r ound seeds and t he second has
gr een wr inkled seeds. All of t he F
1
plant s yield yellow r ound seeds. When you self
fer t ilize t he F
1
t he F
2
gener at ion yields a mixt ur e of yellow r ound, yellow wr inkled,
gr een r ound and gr een wr inkled seeds. What does t his t ell you about t he alleles for
seed color and shape?
(a) t he r ecessive alleles ar e always expr essed
(b) t he alleles ar e on differ ent chr omosomes
(c) t he t wo alleles for each char act er segr egat e dur ing gamet e pr oduct ion
(d) bot h genes a r e on t he sa me chr omosome
11. You cr oss a t r ue-br eeding r ed-flower ed snapdr agon wit h a t r ue-br eeding whit e-flower ed
one. All of t he F
1
ar e pink. What does t his say about t he par ent t r ait s?
(a) r ed and whit e ar e codominant
(b) r ed is dominant
(c) bot h r ed and whit e ar e r ecessive
(d) r ed and whit e show incomplet e dominance
12. While on a field t r ip in t he jungle you find a new species of mouse. You cat ch a pair
and t ake t hem back t o t he lab. In mice, black coat color , B, is dominant t o br own b,
yet t he female mouse gives r ise t o a lar ge lit t er in which 9 of t he offspr ing wer e
black, 3 wer e br own and 4 wer e whit e. You conclude t hat
(a) a new mut at ion has occur r ed in t he mice
(b) t his is an example of polygenic inher it ance
(c) t her e must be an epist at ic int er act ion influencing coat color
(d) t he coat color alleles ar e codominant
13. A new br eed of domest ic cat , t he Indian Cur l Cat , has unusual cur led-back ear s.
When t he owner s of Shulamit h, t he foundat ion cat fr om which t he br eed ar ose, cr ossed
her wit h a nor mal st r aight -ear ed domest ic cat in each of her lit t er s r oughly half of
t he kit t ens had cur led ear s. When bot h par ent s ar e cur l cat s, all t he kit t ens have
cur led ear s. What does t his t ell you about t he cur led-ear t r ait ?
(a) cur led ear s and st r aight ear s ar e codominant t r ait s
(b) cur led ear s and st r aight ear s ar e show incomplet e dominance
(c) cur led ear s ar e dominant
(d) cur led ear s ar e r ecessive
Practice Test Paper–Genetics 225
14. J ohn and J esica ar e planning a family, but since each has a br ot her who has sickle
cell anemia, t hey ar e concer ned t hat t heir childr en may develop sickle-cell disease.
Neit her J ohn, J ane nor t heir r espect ive par ent s have t he disease. They consult a
genet ic counselor who t ells t hem
(a) t her e is ver y lit t le chance t hat any of t heir childr en will have sickle-cell disease
(b) t hat all of t heir childr en will have sickle-cell disease
(c) t hat one out of four of t heir childr en could be expect ed t o have sickle cell-disease
(d) t hat it s possible t hat none of t heir childr en will have t he disease but blood t est s
on t hem bot h will be r equir ed t o make sur e
15. Why is sickle cell disease so called?
(a) because it makes people sick
(b) it s named aft er a special t ype of whit e blood cell
(c) pH changes in t he blood cells make t hem collapse int o a sickle shape
(d) because it s caused by an infect ious micr oor ganism t hat has sickle shaped cells
16. In people wit h sickle cell disease t he r ed blood cells br eakdown, clump, and clog t he
blood vessels. The br oken cells accumulat e in t he spleen. Among ot her t hings t his
leads t o physical weakness, hear t failur e, pain, br ain damage and spleen damage.
Affect ed individuals become par alyzed and can develop r heumat ism, pneumonia and
ot her diseases and kidney failur e. This is an example of
(a) t he polygenic nat ur e of sickle cell disease
(b) t he pleiot r opic effect s of t he sickle cell allele
(c) an epist at ic int er act ion bet ween t he sickle cell allele and a pr ot eolyt ic enzyme
gene
(d) infect ious or ganisms act ing on t he sickle cell allele
17. Hear t disease, diabet es, cancer , alcoholism and many ment al illnesses can best be
descr ibed as:
(a) sympt oms of a bad life-st yle
(b) infect ious diseases caused by micr oor ganisms
(c) mult ifact or ial disor der s wit h a possible polygenic component
(d) all sympt oms of Hunt ingdon’s disease
18. The genet ic disease cyst ic fibr osis is caused by a defect ive allele t hat
(a) pr oduces a dysfunct ional enzyme t hat fails t o br eak down br ain lipids.
(b) causes hemoglobin molecules t o collapse.
(c) pr oduces a defect ive chlor ine-channel membr ane t r anspor t pr ot ein.
(d) pr oduces a neur ot oxin
19. Hunt ingt on’s disease is an example of a genet ic disor der caused by
(a) a lat e-act ing let hal dominant allele
(b) a non-let hal dominant allele
(c) a lat e act ing r ecessive allele
(d) homozygous r ecessive alleles
226 CSIR-NET Life Sciences
20. Which of t he following is a for m of sexual r epr oduct ion?
(a) budding (b) fission
(c) her maphr odit ism (d) r egener at ion
21. The most common phenot ype in a nat ur al populat ion is r efer r ed t o as t he
(a) genot ype (b) wild t ype
(c) aut osome (d) mut ant phenot ype
22. Human males ar e much mor e likely t o be have hemophilia (a failur e of blood t o clot
pr oper ly) t han human females. This is t he case because
(a) hemophilia is a cont agious disease t o which males ar e mor e suscept ible
(b) t he gene for hemophilia is car r ied on t he Y chr omosome
(c) hemophilia is car r ied on t he aut osomes
(d) t he gene for hemophilia is sex-linked
23. In a par t icular species of mammal black hair (B) is dominant t o gr een hair (b) and r ed
eyes (R) ar e dominant t o whit e eyes (r ). If a BbRr individual is mat ed wit h a bbr r
individual t he expect ed phenot ypic r at io of t he offspr ing is 1 black-r ed : 1 black-whit e:
1 gr een-r ed : 1 gr een-whit e. However , when you mat e t hese individuals you find t hat
t he phenot ypic r at io of t he offspr ing is 6 black-r ed : 1 black-whit e : 1 gr een-r ed : 6
gr een-whit e. What could account for t his differ ence?
(a) The genes for hair color and t he genes for eye color ar e car r ied on differ ent
chr omosomes
(b) The expect ed r esult s did not t ake genet ic r ecombinat ion int o account
(c) The genes for hair color and eye color ar e linked
(d) The genes for hair color and eye color show dependent assor t ment
24. In t he pr oblem no. 23 t he obser ved F
2
gener at ion dist r ibut ion of offspr ing was: black-
r ed 1,070; black-whit e 177; gr een-r ed 180; gr een-whit e1072. Based on t his dat a, what
is t he r ecombinat ion fr equency ?
(a) 30 per cent (b) 7 per cent
(c) 17 per cent (d) 14 per cent
25. How many map unit s is a r ecombinat ion fr equency of 5 per cent equal t o?
(a) 2.5 cent imor gans (b) 10 cent imor gans
(c) 5 cent ist ur t evant s (d) 5 cent imor gans
26. A linkage map
(a) or der s genes on a chr omosome based on r ecombinat ion fr equencies
(b) ca n only be const r uct ed for sex chr omosomes
(c) or der s genes on a chr omosome based on t heir locat ion wit h r espect t o a st ained
band
(d) shows t he act ual or der ing and spacing of genes on a chr omosome
27. A male bee is
(a) X Y (b) diploid
(c) hapliod (d) Z W
Practice Test Paper–Genetics 227
28. What is t he pr obabilit y t hat a male will inher it an X-linked r ecessive gene fr om his
fat her ?
(a) 0 (b) 25 per cent
(c) 50 per cent (d) 75 per cent
29. Duchenne muscular dyst r ophy is caused by a sex-linked r ecessive allele. It s vict ims
ar e almost invar iably boys, who usually die befor e t he age of 20. Why is t his disor der
almost never seen in gir ls?
(a) Sex-linked t r ait s ar e never seen in gir ls
(b) The allele is car r ied on t he Y chr omosome
(c) Nondisjunct ion occur s in males but not in females
(d) In or der t o expr ess an X-linked r ecessive, a female must have t wo copies of t he
gene
30. Which of t he following human genet ic disor der s is sex linked?
(a) hemophilia (b) PKU
(c) cyst ic fibr osis (d) achondr oplasia
31. A genet ic defect in humans r esult s in t he absence of sweat glands in t he skin. Some
men have t his defect all over t heir bodies, but in women it is usually expr essed in a
peculiar way. A woman wit h t his defect t ypically has small pat ches of skin wit h sweat
glands and ot her pat ches wher e sweat glands ar e lacking. This pat t er n suggest s t he
phenot ypic effect of
(a) a mut at ion
(b) chr omosome inact ivat ion
(c) RNA splicing
(d) an oper on
32. Which of t he following is cor r ect wit h r egar d t o aneuploidy?
(a) inver sion
(b) 2n + 1
(c) All aneuploid individuals die befor e bir t h
(d) 4n
33. I f a fr a gment of a chr omosome br ea ks off a nd t hen r ea t t a ches t o t he or igina l
chr omosome but in t he r ever se dir ect ion, t he r esult ing chr omosomal abnor malit y is
called
(a) a delet ion (b) an inver sion
(c) a t r anslocat ion (d) a nondisjunct ion
34. Why ar e individuals wit h an ext r a chr omosome 21, which causes Down syndr ome,
mor e numer ous t han individuals wit h an ext r a chr omosome 3 or chr omosome 16?
(a) Ther e ar e pr obably mor e genes on chr omosome 21 t han on t he ot her s
(b) Chr omosome 21 is a sex chr omosome and 3 and 16 ar e not
(c) Down syndr ome is not mor e common, just mor e ser ious
(d) Ext r a copies of t he ot her chr omosomes ar e pr obably fat al
228 CSIR-NET Life Sciences
35. Humans have 23 pair s of chr omosomes, while our closest r elat ives, chimpanzees,
have 24. Chr omosome st udies indicat e t hat at some point ear ly in human evolut ion,
t wo chr omosomes simult aneously br oke int o a lar ge por t ion and a small por t ion. The
lar ge par t s combined t o for m a lar ge chr omosome, and t he small par t s combined t o
for m a much smaller chr omosome (which was subsequent ly lost ). This impor t ant
chr omosomal change could best be descr ibed as
(a) nondisjunct ion followed by delet ion
(b) t r anslocat ion followed by delet ion
(c) duplicat ion followed by delet ion
(d) t r anslocat ion followed by inver sion
36. Each cell in an individual wit h Down syndr ome cont ains ____ chr omosomes.
(a) 47 (b) 22
(c) 24 (d) 45
37. Disor der s involving unusual number s of sex chr omosomes show t hat maleness is
caused by t he
(a) pr esence of a n X chr omosome
(b) pr esence of a Y chr omosome
(c) absence of an X chr omosome
(d) a bsence of a Y chr omosome
38. A par t icular allele can have differ ent effect s if it was inher it ed fr om a male r at her
t han a female. This phenomenon is known as
(a) ext r anuclear inher it ance
(b) genome impr int ing
(c) sex-linkage
(d) Pr ader -Willi syndr ome
39. Human mit ochondr ia
(a) ar e inher it ed as an X-linked t r ait
(b) ar e all inher it ed fr om t he fat her
(c) have linear DNA
(d) ar e all inher it ed fr om t he mot her
40. Bot h chlor oplast s and mit ochondr ia
(a) ar e found wit hin t he nucleus
(b) have linear DNA
(c) car r y ext r anuclear genes
(d) display a Mendelian pat t er n of inher it ance
41. Who demonst r at ed t hat genes ar e locat ed on chr omosomes?
(a) Mor gan (b) Meselson and St ahl
(c) Chargaff (d) Fr anklin
Practice Test Paper–Genetics 229
42. In Gr iffit h’s exper iment s, a har mless var iant of S . pneumoniae became pat hogenic
when mixed wit h a heat -killed pat hogenic var iant as a r esult of
(a) conjugat ion (b) t r ansduct ion
(c) mut at ion (d) t r ansfor mat ion
43. In an impor t ant exper iment , bact er iophages wer e allowed t o infect bact er ia. In t he
fir st t r ial, t he phages used cont ained r adioact ive DNA, and r adioact ivit y was det ect ed
in t he bact er ia. Next , ot her phages cont aining r adioact ive pr ot ein wer e allowed t o
infect ba ct er ia , a nd no r a dioa ct ivit y wa s det ect ed in t he ba ct er ia . When t he
exper iment er s compar ed t he r esult s of t hese t wo t r ials, t hey concluded t hat
(a) genes ar e made of DNA
(b) bact er iophages can infect bact er ia
(c) DNA is made of nucleot ides
(d) genes car r y infor mat ion for making pr ot eins
44. A genet icist r aised a cr op of T2 bact er iophages in a medium cont aining r adioact ive
phosphor us, so t hat t he DNA of t he bact er iophages was labeled wit h r adioact ivit y.
The labeled phages wer e t hen allowed t o infect nonr adioact ive bact er ia. In a few
hour s, t hese bact er ia bur st open, r eleasing many bact er iophages. Some of t hese phages
cont ained labeled
(a) DNA
(b) RNA
(c) pr ot ein
(d) DNA and pr ot ein only
45. Scient ist s have discover ed how t o put t oget her a bact er iophage wit h t he pr ot ein coat
of phage T2 and t he DNA of phage T4. If t his composit e phage wer e allowed t o infect
a bact er ium, t he phages pr oduced in t he host cell would have
(a) t he pr ot ein of T2 and t he DNA of T4
(b) t he pr ot ein of T4 and t he DNA of T2
(c) t he pr ot ein and DNA of T2
(d) t he pr ot ein and DNA of T4
46. Char gaff found t hat for DNA
(a) t he r at io of A t o C is close t o 1:1 and t he r at io of G t o T is close t o 1:1
(b) t he r at io of A t o T is close t o 1:1 and t he r at io of G t o C is close t o 1:1
(c) t he r at io of A t o G is close t o 1:1 and t he r at io of T t o C is close t o 1:1
(d) A + T = G + C
47. The X-r ay diffr act ion st udies conduct ed by ______ wer e key t o t he discover y of t he
st r uct ur e of DNA.
(a) McClint ock (b) Fr anklin
(c) Meselson and St ahl (d) Chargaff
230 CSIR-NET Life Sciences
48. Which of t he following is not t r ue of DNA?
(a) A pair s wit h T and G pair s wit h C
(b) Nit r ogen bases ar e 0.34 nm apar t on a DNA st r and
(c) The double helix is 2.0 nm wide
(d) The double helix is 3.4 nm wide
49. Which of t he following is cor r ect ?
(a) A for ms 2 hydr ogen bonds wit h G; T for ms 3 hydr ogen bonds wit h C
(b) A for ms 3 hydr ogen bonds wit h T; G for ms 2 hydr ogen bonds wit h C
(c) A for ms 2 covalent bonds wit h T; G for ms 3 covalent bonds wit h C
(d) A for ms 2 hydr ogen bonds wit h T; G for ms 3 hydr ogen bonds wit h C
50. Which of t he following is not needed for DNA r eplicat ion?
(a) r ibosomes (b) DNA
(c) nucleot ides (d) enzymes
51. If r ecipr ocal cr oss t o not yield equal r esult it suggest t hat char act er s ar e-
(a) X-linked (b) Aut osomal
(c) Ext r a-chr omosomal (d) None
52. The differ ence which dist inguish pr okar yot ic cell fr om eukar yot ic is-
(a) ER
(b) Mesosome
(c) Nuclear Membr ane
(d) Plasma membr ane
53. Dur ing cr ossing over , exchange of genet ic mat er ial t akes place bet ween
(a) Two chr omat ids
(b) Two chr omosomes
(c) t he non-sist er chr omat ids of t he pair ed chr omosomes
(d) Two sist er chr omat ids of each homologue
54. The segr egat ion of Mendelian fact or s t akes place dur ing-
(a) Meiosis I (b) Meiosis II
(c) Mit osis (d) Int er phase
55. Dur ing Mit osis suddenly t he chr omosomes st ar t s moving t owar d t he opposit e poles
dur ing-
(a) Pr ophase (b) Met aphase
(c) Anaphase (d) Telophase
56. Ext r a nuclear genet ic mat er ial is found in-
(a) Ribosome (b) ER
(c) Chlor oplast (d) Cent r iole
Practice Test Paper–Genetics 231
57. Dosage Compensat ion in case of human is achieved by-
(a) Hyper act ivat ion of X Chr omosome
(b) Hyper act ivat ion of Y chr omosome
(c) Het er ochr omat izat ion of X chr omosome
(d) Het er ochr omat izat ion of Y chr omosome
58. Ext r a nuclear inher it ance is due t o which or ganelle-
(a) Ribosome (b) Cent r iole
(c) Plast id (d) Nucleus
59. The int r oduced cells in a t issue cult ur e ar e made t o divide and for m a mass of
undiffer ent iat ed t issue cells called callus by
(a) adjust ing t he r at io of auxin-cyt okinin
(b) t r ansfer r ing t he plant let s t o pot s in nat ur al envir onment
(c) keeping t he inoculat ed vessels at a desir ed const ant t emper at ur e in an incubat or
(d) solidifying t he enr iched miner al medium wit h agar
60. The fir st st ep in t he t echnique of pr ot oplasmic fusion is t he
(a) hybr idizat ion
(b) collect ion of somat ic cells
(c) isolat ion of pr ot oplast s
(d) select ion and isolat ion of somat ic cells
61. The acr osome of t he sper m is for med fr om t he
(a) mit ochondr ia (b) cent r osome
(c) lysomome (d) golgi bodies
62. A t echnician want ed t o make ant ibody specific for mouse IgM. Accor dingly he inject ed
a r abit wit h pur ified mouse IgM and obt ained an an ant iser um t hat r eact ed st r ongly
wit h mouse IgG. Unfor t unat ely, however ant iser um was also found t o r eact wit h
ot her mouse Ig classes. Such r esult would be obt ained if ant iser um cont ained ant ibodies
dir ect ed against -
(a) The var iable r egion of heavy chain
(b) The const ant r egion of heavy chain
(c) The Fc por t ion of Ig molecule
(d) The light chain of Ig molecule
63. The aver age lengt h of gene is –
(a) 500 bp (b) 1000bp
(c) 2000bp (d) 5000bp
64. Aleur one layer is out er most layer of endosper m in cer eals. It helps in-
(a) Pr ot ect ion of endosper m
(b) Gr owt h of endosper m
(c) Mobilizat ion of r eser ve food in endosper m
(d) Accumulat ion of r eser ve food in endosper m
232 CSIR-NET Life Sciences
65. Genet ically engineer ed male st er ile cr ops plant s have been pr oduced by inser t ing-
(a) Vir al coat pr ot ein gene (b) Chit inase gene
(c) Bar nase gene (d) Opaque Z-gene
66. How many mit ot ic division ar e r equir ed t o pr oduce 12 pollen gr ains in cyper aceae
family-
(a) 2 (b) 6
(c) 3 (d) 12
67. Which plant is used as model syst em t o st udy development al genet ics of zygot ic
embr yogenesis-
(a) Dacus carot a
(b) Arabidposis thaliana
(c) Zea mays
(d) Nicot iana t obacum
68. Your fr iend has just r et ur ned fr om t he exam hall looking ver y ner vous. He was
asked t o choose a met hod for separ at ing 3 amino acids differ ing in t heir polar side
chains. This met hod he choose was paper elect r ophor esis-
(a) You t ell him he choose t he r ight t echnique since t his is a met hod of choice for
separ at ing amino acids wit h differ ent polar it ies
(b) You t ell him he chooses wr ong t echnique r elies on net char ge
(c) You t ell him he is wr ong because t his t echnique also depends on how st r ongly
t he amino acids bind t o paper
(d) You t ell him he is safe as long as he chooses a separ at ion t echnique used for
amino acids
69. Pr ot ein folding is mainly dr iven by all of t he following except -
(a) Hydr ophobic int er act ions (b) Hydr ogen bonds
(c) Covalent bonds (d) Elect r ost at ic at t r act ions
70. How many Bar r bodies would be pr esent in t he WBC of an individual wit h 49 XXXYY
par at ype-
(a) 1 (b) 3
(c) 2 (d) 5
71. An open r eading fr ame is one t hat has-
(a) No st ar t and st op codon (b) A st ar t & st op codon
(c) No st ar t but st op codon (d) A st ar t but no st op codon
72. When t he human genome dr aft sequence was r eleased, which was least expect ed-
(a) The lar ge amount of r epet it ive DNA
(b) The size of t ot al genome
(c) The size of individual chr omosomes
(d) The small number of pr ot ein coding genes
Practice Test Paper–Genetics 233
73. In Sanger ’s Met hod of DNA sequencing, t he gr owing DNA chains ar e t er minat ed
because-
(a) DNA polymer ase is not ver y pr ocessive
(b) A r adioact ive nucleot ide is incor por at ed
(c) The subst r at es become limit at ion
(d) A phosphodiest er bond can not be made
74. A r ecessive mut at ion is t hat -
(a) Not expr essed
(b) Expr essed only when het er ozygous
(c) Expr essed only when homozygous or hemizygous
(d) Eliminat ed by nat ur al select ion
75. Cat alyt ic ant ibodies funct ion as enzymes on t he pr inciple of-
(a) Enzymat ic conver sion of ant ibodies
(b) St abilizing t r ansit ion st at e analogue of subst r at es
(c) Ant igen ant ibody affinit y
(d) Monoclonal ant ibodies wit h chemical
76. In a Sephadex gel filt er at ion column, a mixt ur e of albumin, lysozyme and t hymidine
was loaded. In what sequence t hese will be elut ed fr om t he column-
(a) Albumin > Lysozyme > Thymidine
(b) Lysozyme > t hymidine > Albumin
(c) Thymidine > albumin > Lysozyme
(d) Thymidine > Lysozyme > Albumin
77. Animal vir uses cannot be seen under phase cont r ast micr oscope because-
(a) They do not have a defined mor phology
(b) They ar e t oo small t o be seen under micr oscope
(c) They do not have any envelop t hat make t hem t r anspar ent under micr oscope
(d) Since t hey have no color , t hey ar e not visible
78. You have homogenized plant t issue and would like t o separ at e chlor oplast fr om nuclei.
Which of t he following met hods would be most suit able-
(a) PAGE
(b) Equilibr ium densit y gr adient cent r ifugat ion on CsCl gr adient s.
(c) Differ ent ial cent r ifugat ion using sucr ose gr adient s.
(d) Gel filt r at ion.
79. Which is not ant ibact er ial ant ibiot ic-
(a) Tet r acycline (b) St r ept omycin
(c) Nyst anin (d) Nalidixic acid
234 CSIR-NET Life Sciences
80. Degr adat ion of RNA by RNaseA is an example of-
(a) covalent cat alysis
(b) Acid- base cat alysis
(c) Elect r ost at ic cat alysis
(d) Nucleic acid cat alysis
81. Dehydr ogenase enzymes of hexose mono phosphat e shunt pat hway ar e-
(a) NAD specific (b) NADP specific
(c) FAD specific (d) TPP specific
82. Which of t he following biochemical Rx is most commonly ut ilized by living cells t o
pr opagat e int r acellular signals?
(a) Acylat ion (b) Phosphor ylat ion
(c) Met hylat ion (d) Decar boxylat ion
83. Suppr ession mut at ion r esult s in r est or at ion of wild t ype phenot ype. The suppr ession
of mut ant phenot ype is usually br ought about -
(a) By misr eading of mut ant codon and incor por at ion of a cor r ect amino acid
(b) By inser t ion of anat her copy of gene
(c) By r ever t ion of mut at ion t o wild t ype
(d) Any delet ion of mut ant gene
84 A mouse in which one par t icular gene has been r eplaced by it s inact ivat ed for m
gener at ed in vit r o is called-
(a) Tr ansgenic mouse (b) Nude mouse
(c) Knock out mouse (d) Mut ant mouse
85. Which is not a signal t r ansduct ing molecule-
(a) G pr ot ein coupled r ecept or (b) MAP kinase
(c) Pr ot ein kinase C (d) Insulin
86. Fibr onect in is a pr ot ein found in –
(a) RER (b) Ext r acellular mat r ix
(c) SER (d) Nuclear membr ane
87. Aft er t r anslat ion pr ot eins ar e modified in-
(a) Golgi appar at us (b) Lysosome
(c) Cent r osome (d) Ribosome
88. In AIDS t he pr imar y pr oblem is-
(a) B cells ar e not funct ional
(b) Nat ur al killer cells kill aut ulogous cells
(c) Macr ophages ar e not funct ional
(d) T helper ar e not funct ional
Practice Test Paper–Genetics 235
89. Which one of t he following gr oup of pr ot eins will be most conser ved among differ ent
or ganisms-
(a) Met abolism (b) Tr anscr ipt ion
(c) Tr anslat ion (d) Cell signaling
90. The following ar e samples of r epet it ive element s t hat ar e found in a t ypical eukar yot ic
genome-
(a) r -RNA (b) t -RNA
(c) SINES and LINES (d) Micr osat ellit e
The ascending or der in t er ms of number of r epeat s
(a) ABCD (b) DCBA
(c) CBAD (d) ADBC
91. Which sequence ar e best t o evaluat e t he phylogeny of closely r elat ed mammals-
(a) Coding sequences
(b) Ribosomal pr ot eins
(c) SINES and LINES
(d) Cent r omer ic and t elomer ic sequences
92. Leber her edit ar y opt ic neur opat hy is an inher it ed condit ion what causes a loss of
cent r al vision r esult ing fr om a mut at ion in mit ochondr ial DNA. What is t he pr obabilit y
of t he childr en of a man wit h mut at ions in t wo genes and a woman wit h nor mal
mit ochondr ial DNA inher it ing t his disor der as car r ier s?
(a) 0 % (b) 100 %
(c) 50 % (d) 66.67 %
93. A cont ig fr om t he genome sequence of Plasmodium falciparum, wit h a single st ar t
codon wh en t r a n s l a t ed, wa s fou n d t o h a ve h i gh s i mi l a r i t y wi t h t h e en zyme
dihydr ofolat e r educt ase and t hymidylat e synt hase. Which of following st at ement is
cor r ect -
(a) It cont ains domains pr esent in bot h pr ot eins, but neit her in t wo enzymes
(b) It is single bi-funct ional pr ot ein
(c) Bot h t he pr ot ein shar e a common domain
(d) The pr ot ein is unr elat ed t o bot h t he enzymes. Ther e is a pr oblem wit h t he
similar it y sear ch pr ogr am used for t he st udy
94. The consensus sequence of 5' and 3' splice junct ions in eukar yot ic m-RNA cont ains-
(a) GU-GA (b) GU-AG
(c) AG-GU (d) CG-AG
95. Small cyt oplasmic RNA (Sc-RNA) in eukar yot ic cell-
(a) Splice pr imar y t r anscr ipt
(b) Dir ect pr imar y t r anscr ipt
(c) Dir ect pr ot ein Tr affic
(d) Tr anspor t amino acids
236 CSIR-NET Life Sciences
96. In an exper iment involving polymer ase chain r eact ion (PCR) you have st ar t ed wit h
100 ng of pr imer s and 1 ng of genomic DNA. Aft er a number of cycles, t he same
amount of amplified pr oduct was obser ved. Indicat e which of t he following is t he
r eason for obser vat ion-
(a) Enzymes get s inact ivat ed
(b) Limit at ions of pr imer s
(c) Degr adat ion of t emplat e
(d) Subst r at e inhibit ion of enzyme
97. A genet ic cr oss bet ween t wo F
1
-hybr id pea plant s for spher ical seeds will yield what
per cent spher ical-seeded plant s in t he F
2
gener at ion? (spher ical is dominant over
dent ed)
(a) 100% (b) 75%
(c) 50% (d) 25%
98. A genet ic cr oss bet ween t wo F
1
-hybr id pea plant s having yellow seeds will yield what
per cent gr een-seeded plant s in t he F
2
gener at ion? Yellow seeds ar e dominant t o
gr een.
(a) 100% (b) 25%
(c) 50% (d) 75%
99. When t r ue-br eeding t all st em pea plant s ar e cr ossed wit h t r ue-br eeding shor t st em
pea plant s, all of t he _________ plant s, and 3/4 of t he __________ plant s had t all
st ems. Ther efor e, t all st ems ar e dominant .
(a) F
1
, F
2
(b) G
1
, G
2
(c) par ent al, F
2
(d) F
2
, par ent al
100. To ident ify t he genot ype of yellow-seeded pea plant s as eit her homozygous dominant
(YY) or het er ozygous (Yy), you could do a t est cr oss wit h plant s of genot ype _______.
(a) y (b) Y
(c) yy (d) YY
Practice Test Paper-II
1. mRNA can be isolat ed by passing cell lysat e t hr ough a column of oligo (dT)-Cellulose.
The met hod is example of-
(a) Par t it ion chr omat ogr aphy
(b) Ion-exchange chr omat ogr aphy
(c) Affinit y chr omat ogr aphy
(d) Adsor pt ion chr omat ogr aphy
2. Which of t he following is most commonly involved in globular shape of pr ot ein in
aqueous solut ions-
(a) Hydr ogen bonds (b) Disulphide bonds
(c) Salt br idges (d) Hydr ophobic int er act ions
3. How ma n y di ffer en t ga met es ca n be for med by a n or ga n i s m wi t h gen ot ype
AaBbCCddEe ?
(a) 8 (b) 16
(c) 32 (d) 64
4. In many sit uat ions it has been found t hat t he nucleot ide sequences of t wo highly
homologous pr ot eins ar e differ ent . It may be due t o use of-
(a) differ ent amino acids
(b) non-over lapping genes
(c) differ ent st r ands of DNA for encoding gene
(d) synonymous codons due t o degener acy
5. DNA binding pr ot ein t hat pr event s t r anscr ipt ion ar e-
(a) Act ivat or s (b) Oper at or s
(c) Repr essor s (d) Silencer s
6. The pollen t ube dischar ges it s cont ent in-
(a) The egg (b) One of syner gid
(c) The cent r al cell (d) Ant ipodal Cell
7. The spindle fibr es in a mit ot ic cell ar e composed of-
(a) Chr omat in (b) Act in
(c) Nucleopr ot ein (d) Tubulin
8. In het er ozygous cell t he gene of r ecessive allele
(a) Is always t ur ned off
(b) May be expr essed but pr oduct is quickly degr aded
(c) May be expr essed wit hout measur able effect
(d) Is delet ed
9. The fir st st ep in t he init iat ion of pr ot ein synt hesis r equir es
(a) The 30 S r ibosomal subunit (b) ATP
(c) The r ibosomal A sit e (d) Tr anslat ional fact or Tu
238 CSIR-NET Life Sciences
10. House keeping genes ar e-
(a) inducible genes
(b) expr essed only in t umour cells
(c) expr essed in all cells
(d) do not expr ess at all
11. Reduct ion of chr omosome number occur s in-
(a) Mit ot ic anaphase
(b) Anaphase –I of meiosis
(c) Anaphase-II of meiosis
(d) Mit ot ic as well as meiot ic anaphases
12. When a plant of chr omosomal t ype aa pollinat es a plant of t ype AA, what chr omosome
const it ut ion of embr yo and endosper m is expect ed in t he r esult ing seeds-
(a) Diploid zygot e of t ype Aa and Tr iploid endosper m of t ype AAa
(b) Diploid zygot e of t ype aa and Tr iploid endosper m of t ype Aaa
(c) Diploid zygot e of t ype AA and Tr iploid endosper m of t ype AAa
(d) Diploid zygot e of t ype Aa and Tr iploid endosper m of t ype aaa
13. A cult ur e of t et r acycline sensit ive bact er ia was infect ed by a phage t hat is deliver ed
fr om t he lysis of a t et r acycline r esist ant bact er ial st r ain. This r esult s in development
of t et r acycline r esist ant in t he or iginal cult ur e. What phenomenon has occur r ed-
(a) Conjugat ion (b) Recombinat ion
(c) Tr ansfor mat ion (d) Tr ansduct ion
14. If t he molecular mass of amino acid is 150 dalt on, t he molecular weight of it s t r ipept ide
will be-
(a) 450 (b) 486
(c) 504 (d) 414
15. If t he number of het er ozygous pair s involved in a par t icular cr oss is t hr ee, t he
phenot ype r at io obt ained in t heir F
2
gener at ion will be-
(a) 9:3:3:1 (b) 27:9:9:9:6:6:3:3:1
(c) 27:9:9:9:3:3:3:1 (d) 1:2:1:2:4:2:1:2:1
16. Rheumat oid ar t hr it is is a/an-
(a) immunodeficiency disease
(b) sexually t r ansmit t ed disease
(c) insect bor n disease
(d) aut oimmune disease
17. A linkage gr oup cor r esponds t o a-
(a) chr omosome
(b) set of independent ly assor t ing genes
(c) set of independent ly segr egat ing alleles
(d) set of non complement ing alleles
Practice Test Paper–Genetics 239
18. Which of t he following r est r ict ion enzymes pr oduces blunt end DNA fr agment s-
(a) EcoR I (b) EcoR II
(c) EcoR V (d) Bam H1
19. Who discover ed t hat DNA is t he genet ic mat er ial-
(a) Ar t hur Kor nber g (b) J ames Wat son
(c) Ost awald aver y (d) Sever o Ochoa
20. Which of t he common bases (A,C,G,T) if DNA has no oxygen in it s st r uct ur e-
(a) Thymine (b) Adenine
(c) Cyt osine (d) Guanine
21. What is main damaging effect of UV r adiat ions on DNA-
(a) Depur inat ion
(b) For mat ion of t hymine dimmer s
(c) Single st r and br eak
(d) Double st r and br eak
22. In a cr oss bet ween a pur e t all pea plant wit h gr een pods and pur e dwar f pea plant
wit h yellow pods, how many plant s in F
2
gener at ion will be dwar f out of 16 plant s-
(a) 1 (b) 3
(c) 4 (d) 9
23. If par ent s have AB and O blood gr oup, t heir offspr ing could be of-
(a) O gr p only (b) A & B gr p
(c) A, B, O gr p (d) A, B, O and AB
24. Similar set of r egulat or y genes cont r ol development in Ar abidopsis, Dr osophila and
mice. These genes ar e called-
(a) Homologous genes (b) Het er ologous
(c) Homeot ic (d) Or t hologous
25. TATA box and Pr ibnow box ar e component s of-
(a) Oper at or s (b) Pr omot er s
(c) Enhancer s (d) Act ivat or s
26. Lampbr ush chr omosomes occur t hr ough-
(a) diplot ene of meiosis (b) Pr ophase of mit osis
(c) Int er phase (d) Met aphase of meiosis
27. Which of t he following is not associat ed wit h cell cycle-
(a) Cyclins (b) Myosins
(c) CDK (d) DNA polymer ases
28. Segr egat ion of alleles t akes place dur ing t he following st age of meiot ic division-
(a) Met aphase I (b) Anaphase I
(c) Diplot ene (d) Anaphase II
240 CSIR-NET Life Sciences
29. A single cr ossing over bet ween t wo homologous chr omosomes involves-
(a) Two chr omat ids
(b) Thr ee chr omat ids
(c) Four chr omat ids
(d) The cent r omer e of t he chr omosomes
30. Haploids ar e consider ed bet t er genet ic st ock because t hey-
(a) Ar e healt hier t hen diploids
(b) Requir e only half of t he nut r ient s
(c) Easy t o cult ur e
(d) For m homozygous individual on doubling
31. A yellow r ound seeded pea plant is cr ossed wit h gr een and wr inkled pea plant and t he
F1 of t his ar e back cr ossed t o t he homozygous r ecessive par ent s. The pr ogeny will
appear -
(a) 9:3:3:1 (b) 12:3:3:1
(c) 3:3:3:1 (d) 1:1:1:1
32. Which of t he following is t he appr oximat ely size of human genome-
(a) 4 × 10
6
bp (b) 1 × 10
10
bp
(c) 3 × 10
9
bp (d) 5 × 10
5
bp
33. Which of t he following t ype of DNA is t he most conser ved amongst or ganisms-
(a) Mit ochondr ial DNA (b) Chlor oplast DNA
(c) r DNA (d) DNA t hat codes for t -DNA
34. Which of t he following is incor r ect r egar ding char gaff’s r ule-
(a) [A] = [T] (b) [G] =[C]
(c) [A] + [T] = [G] + [C] (d) [A ]+ [G] = [T] + [C]
35. If t ot al concent r at ion of A=T is 56 %. What will be concent r at ion of cyt osine in genome:
(a) 56 (b) 23
(c) 44 (d) 22
36. In t he B for m of DNA, t he pair ed bases ar e planar , par allel t o one anot her and t hey
a r e-
(a) Par allel t o long axis of double helix
(b) Per pendicular t o long axis of double helix
(c) Inclined t o long axis
(d) Inver t ed in r espect t o long axis
37. Among t he Following st agger ed cut is obt ained by-
(a) Alu I (b) Rsa I
(c) Pst I (d) Pvu (II)
38. Which one is t he essent ial feat ur e of r est r ict ion sit es cut by r est r ict ion endonucleases
II-
(a) Palindr omic (b) Wit hin r ecognisat ion sit e
(c) Always met hylat ed (d) All t he above
Practice Test Paper–Genetics 241
39. The genomic size of E. coli is 4.6 X 10
6
bp, if a 6 base pair cut t er is ut ilized t o obt ain
r est r ict ion fr agment s, t hen what will be t ot al number of fr agment s obt ained-
(a) 1.12 × 10
3
(b) 7.66 × 10
5
(c) 25 × 10
4
(d) 3.83 × 10
3
40. Tm (Melt ing t emper at ur e) would be maximum for -
(a) S aricina lutea (b) E. Coli
(c) Dr osophila (d) Human
41. Nat ur al absor pt ion of UV by DNA is due t o nucleot ide base pair s which incr ease on
denat ur at ion. They absor b maximum amount of UV wavelengt h of-
(a) 200 nm (b) 260 nm
(c) 280 nm (d) 355 nm
42. For Nucleic acid hybr idizat ion which condit ion is not essent ial –
(a) The salt conc. must be high (>25 M)
(b) Pr imer s
(c) High t emper at ur e, under cont r ol
(d) Complement ar y st r ands
43. Genet ic var iat ions, in for m of mult iple alleles of many genes, exist in most of nat ur al
popu l a t i on . Su ch gen et i c di ffer en ces bet ween i n di vi du a l s a r e ca l l ed DNA
polymor phism. Such a DNA polymor phism is used as-
(a) DNA mar ker s (b) DNA pr obes
(c) c-DNA (d) Het er oduplex DNA
44. The best DNA mar ker s ut ilized t o differ ent iat e differ ent human individuals (t o
est ablish pat er nit y & diver gence) ar e-
(a) Single Nucleot ide Polymor phism
(b) Rest Fr agment Lengt h Polymor phism
(c) Random Amplified Polymor phic DNA
(d) Simple t andem r epeat s polymor phism
45. Which chemical gr oup ar e pr esent at t he ext r eme 3’ ends of single polynucleot ide
st r and-
(a) Phosphat e gr oup (b) Hydr oxyl gr oup
(c) Oxo gr oup (d) CH
2
OH
46. The t emplat e DNA st r and ut ilized for DNA r eplicat ion is-
(a) 5' -3' dir ect ion (b) 3' -5 dir ect ion
(c) Random (d) In bot h dir ect ion
47. The enzyme coded by allele I
A
t r ansfer ase which adds N-acet yl glucasmine and blood
gr oup is designat ed as A, similar ly I
B
encodes t r ansfer ase which adds galact ose t o
pr ecur sor sugar , similar ly I
O
encodes t r ansfer ase which adds-
(a) No sugar
(b) N-acet yl glucasamine
(c) Galact ose
(d) Bot h galact ose and N acet yl glucasmine
242 CSIR-NET Life Sciences
48. If t wo r ecessive mut at ions ar e alleles of differ ent genes, t hen F
1
pr ogeny ar e wild
t ype, t his is due t o-
(a) Incomplet e dominance (b) Complement at ion
(c) Supplement at ion (d) Co-dominance
49. Sexual differ ent iat ion in Dr osophila is cont r olled by gene called sex let hal (sxl). The
genes sensing t he number of x chr omosomes ar e called as-
(a) Super numer y genes (b) Denomina t or genes
(c) Numer a t or genes (d) Sex sensing genes
50. What would be sex of Dr osophila wit h chr omosome complement 3A + XXYY-
(a) Male (b) Female
(c) Inet er sex (d) Met amale
51. Eukar yot ic gene r egulat ion occur s at t he level of:
(a) post t r anslat ional cont r ol (b) t r anscr ipt ional cont r ol
(c) genomic cont r ol (d) all of t he above
52. Ubiquit in binds t o _____ r esidues, t her efor e t ar get ing pr ot eins for degr adat ion by
_____.
(a) lysine, pr ot easomes (b) ar ginine, lysosomes
(c) ar ginine, pr ot easomes (d) lysine, lysosomes
53. Pr ot ein phosphor yat ion, dephosphor ylat ion and pr ot eolyt ic cleavage ar e examples of:
(a) post t r anslat ional cont r ol
(b) t r anscr ipt ional cont r ol
(c) t r anslat ional cont r ol
(d) cont r ol of RNA pr ocessing
54. St er oid hor mone r ecept or s ar e involved in:
(a) cont r ol of RNA pr ocessing
(b) post t r anslat ional cont r ol
(c) t r anscr ipt ional cont r ol
(d) genomic cont r ol
55. Which of t he following is NOT an example of genomic cont r ol of gene r egulat ion?
(a) alt er nat ive RNA splicing
(b) DNA r ear r angement
(c) gene delet ion
(d) gene amplificat ion
56. Which of t he following is t r ue of homeot ic genes?
(a) t he homeodomain funct ions in binding t o DNA
(b) t hey ser ve as an example of t r anscr ipt ional gene cont r ol
(c) all homeot ic genes cont ain a 180-bp segment called a homeobox
(d) all of t he above ar e cor r ect
Practice Test Paper–Genetics 243
57. ______ is an allost er ic pr ot ein t hat is inact ive unt il it binds t o_____ , t hus act ivat ing
t r anscr ipt ion.
(a) RNA polymer ase, cAMP (b) CRP, ATP
(c) CRP, cAMP (d) RNA polymer ase, ATP
58. The t r p leader sequence cont ains a cont r ol r egion t hat is sensit ive t o t r ypt ophan
levels such t hat it det er mines whet her t r anscr ipt ion will cont inue t o complet ion. The
effect of t his cont r ol element in t he t r p oper on in E. coli is known as:
(a) gene amplificat ion (b) RNA splicing
(c) at t enuat ion (d) r epr ession
59. Hist one st r uct ur e can be alt er ed by:
(a) DNA r ear r angement (b) acet ylat ion
(c) sigma fact or s (d) met hylat ion
60. Which of t he following is t r ue of heat -shock genes?
(a) t hey ar e only found in pr okar yot es
(b) t hey encode for pr ot eins such as pr ogest er one and est r ogen
(c) t hey ar e only found in eukar yot es
(d) t hey ar e known t o r espond t o st r essful condit ions
61. A gene is
(a) t he same t hing as a chr omosome.
(b) t he infor mat ion for making a polypept ide.
(c) made of RNA.
(d) made by a r ibosome.
62. In Eukar yot es, DNA packing affect s gene expr ession by
(a) cont r olling access t o DNA.
(b) posit ioning r elat ed st r uct ur al genes near each ot her .
(c) pr ot ect ing DNA fr om mut at ions.
(d) enhancing r ecombinat ion of genes.
63. Dioxin, pr oduced as a by-pr oduct of var ious indust r ial chemical pr ocesses, is suspect ed
of causing cancer and bir t h defect s in animals and humans. It appar ent ly act s by
ent er ing cells and binding t o pr ot eins, alt er ing t he pat t er n of gene expr ession. The
pr ot eins affect ed by dioxin ar e pr obably
(a) wat er -soluble pr ot eins. (b) DNA polymer ase.
(c) t r anscr ipt ion fact or s. (d) enha ncer s.
64. In humans, t he hor mone t est ost er one ent er s cells and binds t o specific pr ot eins,
which in t ur n bind t o specific sit es on t he cells DNA. These pr ot eins pr obably act t o
(a) help RNA polymer ase t r anscr ibe cer t ain genes.
(b) alt er t he pat t er n of DNA splicing.
(c) inhibit t r anscr ipt ion.
(d) unwind t he DNA for gene t r anscr ipt ion.
244 CSIR-NET Life Sciences
65. It is possible for a cell t o make pr ot eins t hat last for mont hs; hemoglobin in r ed blood
cells is a good example. However , many pr ot eins ar e not t his long-last ing. They may
be degr aded in days or even hour s. Why do cells make pr ot eins wit h such shor t
lifet imes if it is possible t o make t hem last longer ?
(a) Most pr ot eins ar e used only once
(b) Most cells in t he body live only a few days
(c) Cells lack t he r aw mat er ials t o make most of t he pr ot eins t hey need
(d) Only cancer cells, which can keep dividing, cont ain long-last ing pr ot eins.
66. The genes t hat malfunct ion in cancer nor mally
(a) cont r ol RNA t r anscr ipt ion
(b) ar e r esponsible for sex det er minat ion
(c) code for enzymes t hat r epair damaged DNA
(d) ar e not pr esent in most body cells
67. Which of t he following ar e ar r anged in t he cor r ect or der by size, fr om lar gest t o
smallest ?
(a) chr omosome-gene-codon-nucleot ide
(b) nucleot ide-chr omosome-gene-codon
(c) codon-chr omosome-gene-nucleot ide
(d) gene-chr omosome-codon-nucleot ide
68. Imagine an er r or occur r ing dur ing DNA r eplicat ion in a cell, so t hat wher e t her e is
supposed t o be a T in one of t he genes t her e is inst ead a G. What effect will t his
pr obably have on t he cell?
(a) Each of it s kinds of pr ot ein will cont ain an incor r ect amino acid
(b) An amino acid will be missing fr om each of it s kinds of pr ot ein
(c) One of it s kinds of pr ot ein might cont ain an incor r ect amino acid
(d) The amino acid sequence of one of it s kinds of pr ot ein will be complet ely changed
69. How does RNA polymer ase know wher e t o st ar t t r anscr ibing a gene int o mRNA?
(a) Tr ansfer RNA act s t o t r anslat e t he message t o RNA polymer ase
(b) It st ar t s at a cer t ain nucleot ide sequence called a pr omot er
(c) The r ibosome dir ect s it t o t he cor r ect por t ion of t he DNA molecule
(d) It looks for t he AUG st ar t codon
70. All your cells cont ain pr ot o-oncogenes, which can change int o cancer -causing genes.
Why do cells possess such pot ent ial t ime bombs?
(a) Vir uses infect cells wit h pr ot o-oncogenes
(b) Pr ot o-oncogenes ar e genet ic junk and have no known funct ion
(c) Pr ot o-oncogenes ar e unavoidable envir onment al car cinogens
(d) Cells pr oduce pr ot o-oncogenes as a by-pr oduct of mit osis
71. In Eukar yot es, which of t he following mechanisms of gene r egulat ion oper at es aft er
mRNA t r anscr ipt ion but befor e t r anslat ion of mRNA int o pr ot ein?
(a) mRNA splicing and edit ing (b) DNA packing
(c) r epr essor s and act ivat or s (d) pr ot ein degr adat ion
Practice Test Paper–Genetics 245
72. A cell biologist found t hat t wo differ ent pr ot eins wit h lar gely differ ent st r uct ur es
wer e t r anslat ed fr om t wo differ ent mRNAs. These mRNAs, however , wer e t r anscr ibed
fr om t he same gene in t he cell nucleus. Which mechanism below could best account
for t his?
(a) Differ ent syst ems of DNA unpacking could r esult in t wo differ ent mRNAs
(b) A mut at ion might have alt er ed t he gene
(c) Exons fr om t he same gene could be spliced in differ ent ways t o make differ ent
mRNAs
(d) The t wo mRNAs could be t r anscr ibed fr om differ ent oper ons
73. A par t icular ____ car r y t he infor mat ion for making a par t icular polypept ide, but ____
can be used t o make any polypept ide.
(a) gene and r ibosome . . . a t RNA and an mRNA
(b) gene and mRNA . . . a r ibosome and a t RNA
(c) r ibosome and mRNA . . . a gene and a t RNA
(d) gene and t RNA . . . a r ibosome and an mRNA
74 Which of t he following pr ocesses occur s in t he cyt oplasm of a eukar yot ic cell?
(a) DNA r eplicat ion
(b) t r anslat ion
(c) t r anscr ipt ion
(d) DNA r eplicat ion and t r anslat ion
75. The nucleot ide sequence of a DNA codon is GTA. A messenger RNA molecule wit h a
complement ar y codon is t r anscr ibed fr om t he DNA. In t he pr ocess of pr ot ein synt hesis,
a t r ansfer RNA pair s wit h t he mRNA codon. What is t he nucleot ide sequence of t he
t RNA ant icodon?
(a) CAT (b) GUA
(c) CAU (d) GTA
76. The nucleot ide sequence of a DNA codon is ACT. A messenger RNA molecule wit h a
complement ar y codon is t r anscr ibed fr om t he DNA. In t he pr ocess of pr ot ein synt hesis,
a t r ansfer RNA pair s wit h t he mRNA codon. What is t he nucleot ide sequence of t he
t RNA ant icodon?
(a) TGA (b) UGA
(c) TGU (d) ACU
77. Dur ing t he pr ocess of t r anslat ion (polypept ide synt hesis), ____ mat ches an mRNA
codon wit h t he pr oper amino acid.
(a) a r ibosome (b) DNA polymer ase
(c) ATP (d) t r ansfer RNA
78. A sequence of pict ur es of polypept ide synt hesis shows a r ibosome holding t wo t r ansfer
RNAs. One t RNA has a polypept ide chain at t ached t o it ; t he ot her t RNA has a single
amino acid at t ached t o it . What does t he next pict ur e show?
(a) The polypept ide chain moves over and bonds t o t he single amino acid
(b) The amino acid moves over and bonds t o t he polypept ide chain
(c) The t RNA wit h t he polypept ide chain leaves t he r ibosome
(d) A t hir d t RNA wit h an amino acid joins t he pair on t he r ibosome
246 CSIR-NET Life Sciences
79. A genet icist found t hat a par t icular mut at ion had no effect on t he polypept ide coded
by a gene. This mut at ion pr obably involved
(a) delet ion of one nucleot ide
(b) alt er at ion of t he st ar t codon
(c) inser t ion of one nucleot ide
(d) subst it ut ion of one nucleot ide
80. A mut agen is
(a) a gene t hat has been alt er ed by a mut at ion
(b) somet hing t hat causes a mut at ion
(c) an or ganism t hat has been changed by a mut at ion
(d) t he por t ion of a chr omosome alt er ed by a mut at ion
81. Ther e ar e t hought t o be about ____ genes in a human cell.
(a) 30 – 100 (b) 300 – 1,000
(c) 3,000 – 10,000 (d) 30,000 – 50,000
82. Hist ones ar e
(a) mast er genes t hat affect development
(b) gr oups of genes t hat r espond t o envir onment
(c) pr ot eins ar ound which DNA is coiled
(d) por t ions of genes t hat ar e t r anscr ibed
83. Dur ing Int er phase, _____ can be seen wit h a light micr oscope.
(a) nucleosomes (b) int r ons
(c) het er ochr omat in (d) euchr omat in
84. Ther e is about 1,000 t imes as much DNA in a human cell as in an E. coli cell, but only
about 50 t imes as many genes. Why?
(a) A human cell has much mor e noncoding DNA
(b) The DNA packing is much mor e complex in a pr okar yot ic cell
(c) Most of t he genes in a human cell ar e t ur ned off
(d) E. coli ar e less able t o r espond t o t heir envir onment t han humans. Mor eover ,
t his r esponse confuses cause and effect
85. The differ ence bet ween t andemly r epet it ive and int er sper sed r epet it ive DNA is t hat
(a) int er sper sed DNA is also r efer r ed t o as sat ellit e DNA
(b) int er sper sed r epet it ive DNA is found t hr oughout t he genome.
(c) most t andemly r epet it ive DNA ar e t r ansposons
(d) most int er sper sed r epet it ive DNA is at t he t elomer es
86. Mult igene families ar ise as a r esult of
(a) t r ansfor mat ion
(b) er r or s dur ing DNA r eplicat ion and r ecombinat ion
(c) RNA splicing
(d) pr ot ein degr adat ion
Practice Test Paper–Genetics 247
87. Ret r ot r ansposons differ fr om ot her t r ansposons in t hat
(a) r et r ot r ansposons have lost t he abilit y t o move about a genome
(b) r et r ot r ansposons ar e likely t o be t he r emains of a vir al infect ion
(c) r et r ot r ansposons have r et ained t he abilit y t o move about a genome, an abilit y
t hat has been lost by ot her t r ansposons
(d) r et r ot r ansposons move via an RNA t r anscr ipt , wher eas ot her t r ansposons do
not
88. Your muscle and bone cells ar e differ ent because
(a) t hey cont ain differ ent set s of genes
(b) t hey ar e differ ent iat ed
(c) t hey cont ain differ ent oper ons
(d) differ ent genes ar e swit ched on and off in each t ype of cell
89. Gene expr ession in animals seems t o be r egulat ed lar gely by
(a) cont r olling gene packing and unpacking
(b) cont r olling t he t r anscr ipt ion of genes int o mRNA
(c) cont r olling t he t r anslat ion of mRNA int o pr ot ein
(d) select ively eliminat ing cer t ain genes fr om t he genome
90. The cont r ol of gene expr ession is mor e complex in mult icellular eukar yot es t han in
pr okar yot es because
(a) eukar yot ic cells ar e much smaller
(b) in a mult icellular eukar yot e, differ ent cells ar e specialized for differ ent funct ions
(c) pr okar yot es ar e r est r ict ed t o st able envir onment s
(d) eukar yot es have fewer nucleot ide, so each nucleot ide sequence must do sever al
jobs
91. Which of t he following would be most likely t o lead t o cancer ?
(a) mult iplicat ion of a pr ot o-oncogene and inact ivat ion of a t umor -suppr essor gene
(b) hyper act ivit y of a pr ot o-oncogene and act ivat ion of a t umor -suppr essor gene
(c) failur e of a pr ot o-oncogene t o pr oduce a pr ot ein and mult iplicat ion of a t umor -
suppr essor gene
(d) t he failur e of bot h a pr ot o-oncogene and a t umor -suppr essor gene t o pr oduce
pr ot eins
92. Your bone cells, muscle cells, and skin cells look differ ent because
(a) differ ent kinds of genes ar e pr esent in each kind of cell
(b) t hey ar e pr esent in differ ent or gans
(c) differ ent genes ar e act ive in each kind of cell
(d) t hey cont ain differ ent number s of genes
93. Linkage gr oups ar e equivalent t o haploid set of chr omosomes, if male but t er fly has
12 linkage gr oups, t hen female will have-
(a) 12 (b) 11
(c) 13 (d) 6
248 CSIR-NET Life Sciences
94. Maximum possible r ecombinat ion fr equency is:
(a) 25 % (b) 50 %
(c) 75 % (d) 100 %
95. The linkage of genes in chr omosomes can be r epr esent ed in for m of-
(a) genet ic maps (b) Linkage maps
(c) Chr omosome map (d) All
96. Physically 1 map unit on linkage maps can be defined as lengt h of chr omosome in
which, aver age cr ossover for med dur ing 50 cells under going meiosis is-
(a) 1 (b) 25
(c) 50 (d) 100
97. Assuming equal sex r at io, what is pr obabilit y t hat a sib ship of four childr en consist s
ent ir ely of boys-
(a) 25 % (b) 12. 5 %
(c) 6.25 % (d) 3.125 %
98. The spindle fibr es at t ach t o each chr omosome in t he r egion t echnically known as-
(a) Cent r omer e (b) Cent r iole
(c) Kinet ochoer e (d) Ast r als
99. Which among t he following do not have DNA-
(a) Kinet oplast (b) Cent r iole
(c) Dict yosomes (d) Chindr iosomes
100. Dur ing meiosis t he cent r omer ic division t akes place dur ing-
(a) Pr opha se I (b) Anaphase I
(c) Pr ophase II (d) Anaphase II
5
Evolutionary Biology
EARLY EARTH AND THE ORIGIN OF LIFE
ORIGIN OF EARTH AND PRE-BIOTIC ENVIRONMENT
• Our planet and our solar syst em has a hist or y of about 4.55 t o 4.60 Ga (Ga = billion
year s). Our solar syst em or iginat ed LONG aft er t he or igin of t he univer se in t he Bi g
Ba n g about 10 t o 15 billion year s ago. Bi g Ba n g is a t heor et ical explosion of t hick
concent r at ed cosmic mat t er t hat occur r ed 10,000-20,000 million year s ago t o for m
our univer se and 100,000 galaxies. It was suggest ed by Ab b e Le ma i t r e (1931).
• All mat t er for med dur ing t he Big Bang consist ed of t he element hydr ogen. The
hydr ogen at oms had t o under go nuclear r eact ions in st ar s, and t he st ar s had t o explode
as super novas, befor e heavier element s t han hydr ogen (such as car bon, t he main
building block of life on Ear t h) came int o exist ence. Our Sun did t hus not or iginat e
st r aight at t he or igin of t he univer se.
• Sun, ear t h and ot her st ar s ar e for med fr om Nebula (cosmic dust s and clouds of gases).
Nebular hypot hesis of Ka n t is for or igin of solar syst em. The or igin of our univer se
is 10-20 billion year s (10,000-20,000 million year s or 10
4
t o 20
4
million year s) old. The
univer se has 1 lakh million (100000 million) galaxies and 10000 million st ar s in t he
milky way.
• The Sun was not as hot as it is t oday. The amount of solar r adiat ion r eaching t he
Ear t h was only 70-80% of t hat of t oday, because nuclear r eact ions in t he Sun wer e in
an ear lier st age – we know t hat fr om obser vat ions on ot her st ar s.
• At fir st t her e was no Moon. The chemical composit ion of Moon r ocks and t he way in
which t he Moon moves t oget her wit h t he Ear t h ar ound t he Sun indicat e t hat t he
Moon was for med by t he collision of a Mar s-size ast er oid wit h ear ly Ear t h, pr obably
at some t ime dur ing t he big met eor bombar dment t hat ended about 4 billion year s
ago (4 Ga). The mat er ial fr om t he cor e of t he big ast er oid was added t o Ear t h, it s
250 CSIR-NET Life Sciences
mant le moved on and for med t he Moon. The Moon was closer t o Ear t h aft er it
or iginat ed, so t he t ides wer e much mor e pr onounced t han t oday’s t ides.
• The Ear t h’s r ot at ion r at e was fast er because t idal fr ict ion r esult ing fr om t he pr esence
of t he Moon had not yet slowed it down. Days may have been bet ween 8 and 14 hour s
long. Gr owt h r ings in Paleozoic cor als shows t hat even as r ecent ly as 450 million
year s ago t her e wer e mor e t han 400 days in a year (a year is t he t ime t hat it t akes
t he Ear t h t o cir cle t he Sun). Ear t h in t he beginning was a hot spinning ball wit h a
t emper at ur e of 5273-6273 Kelvin (5000-6000 °C). The cont inent s wer e pr obably smaller
t han t oday’s, wit h mor e ocean sur face, especially mor e t han 3 billion year s ago.
• The at mospher e had a ver y differ ent composit ion. Our at mospher e at t he moment
consist s for about 80% of dinit r ogen gas (N
2
) and 20% oxygen gas (O
2
), wit h t r aces of
ot her gases. CO
2
(car bon dioxide) at t he moment makes up about 360 par t s per million
of t he at mospher e (much less t han 1%, which is one par t per hundr ed). Befor e t he
indust r ial r evolut ion at mospher ic CO
2
levels wer e about 280 ppm.
• The ear ly Ear t h’s at mospher e was differ ent : t her e was no fr ee oxygen gas. The pr imar y
at mospher e of t he Ear t h, inher it ed fr om t he swir ling cloud of gas fr om our solar
syst em for med, would have been dominat ed by hydr ogen gas, H
2
, wit h ammonia,
NH
3
, and met hane, CH
4
. At mospher e was r educing and called at mospher e I. Pr esent
at mospher e is oxidizing and is called at mospher e II and has about 21% oxygen.
• Hydr ogen gas is t oo light for a planet wit h t he size of Ear t h t o hold by it s gr avit y, and
t he pr imar y at mospher e would have been blown away by t he violent r adiat ion of t he
Sun in it s so-called T-t aur i st age. A secondar y at mospher e would t hen be supplied by
out gassing fr om t he Ear t h’s int er ior ; eit her r apidly (big - t heor y) or mor e gr adual.
This secondar y at mospher e consist ed of dominant ly CO
2
(car bon dioxide), wit h some
N
2
(dinit r ogen gas), H
2
O (wat er vapor ), minor CO (car bon monoxide), SO
2
(sulfur
dioxide), and H
2
S (hydr ogen sulfide).
• Bi op oi e s i s is t he st udy of or igin of life on ear t h. Cos mol ogy is t he st udy of univer se
(cosmos).
MAJOR EPISODES IN THE HISTORY OF LIFE: A PREVIEW
• Life or iginat ed in Ar chaeozoic er a but fir st evidence of life is fr om Pr ecambr ian per iod
of Pr ot er ozole er a in ocean about 3.6-4.2 billion year s ago. Scient ist s have found
isot opes of car bon in 3.8 billion year old r ocks in Gr eenland.
• Oldest r ecor d of fossils is 3200-3500 (3.2 t o 3.5 billion) million year s old. The oldest
r ecor ded fossil is fr om t he banded domes of calcar eous sediment s (St r omat olit es) in
Zimbawe (Rhodesia). It is 2.9 billion year s old. Age of fossils is det er mined eit her by
14
C dat ing t echnique or by amount of lead in a r ock.
• Fossil evidence suggest s t hat pr okar yot es appear ed at least 2 billion year s befor e t he
oldest eukar yot es
• Two dist inct gr oups of pr okar yot es, Bact er ia and Ar chaea, diver ged ear ly, bet ween 2
t o 3 billion year s ago. Phot osynt het ic bact er ia st ar t ed t he pr oduct ion of oxygen about
2.5 billion year s ago, set t ing t he st age for aer obic life.
• Eukar yot es emer ged some 2 billion year s ago. St r ong evidence suppor t s t he hypot hesis
t hat eukar yot ic cells evolved fr om a symbiot ic communit y of pr okar yot es.
Evolutionary Biology 251
• Plant s, fungi, and animals ar ose fr om dist inct gr oups of unicellular eukar yot es dur ing
he Pr ecambr ian. Plant s evolved fr om gr een algae. Fungi and animals ar ose fr om
differ ent gr oups of het er ot r ophic unicells. Based on molecular evidence, fungi ar e
mor e closely r elat ed t o animals t han t hey ar e t o plant s.
• The oldest fossils of animals ar e t hose of soft -bodied inver t ebr at es fr om about 700
million year s ago. The basic body plans of most of t he moder n animal phyla pr obably
ar ose in t he lat e Pr ecambr ian.
• The t r ansit ion fr om t he aquat ic envir onment t o land was a pivot al point in t he hist or y
of life. The fir st t er r est r ial colonizat ion was by plant s and fungi some 475 million
year s ago (Paleozoic); t he move may have depended upon a beneficial associat ion
bet ween t he t wo gr oups. The t r ansfor mat ion of t he landscape by plant s cr eat ed new
oppor t unit ies for all for ms of life.
• Or igin of life and evolut ion of life ar e independent pr ocesses. Or igin of life is a chemical
pr ocess and evolut ion of life is a biological pr ocess.
PREBIOTIC CHEMICAL EVOLUTION AND THE ORIGINS OF LIFE
• The fir st life wa s a na er obic ch e moh e t e r ot r op h i c ma r i n e p r ok a r yot i c. Fir st
phot oaut ot r ophs wer e anoxygenic and anaer obic chemoaut ot r ophos. Ribonucleot ides
appear ed pr ior t o deoxyr ibonucleot ides.
• Appear ance of nucleopr ot eins gave t he fir st sign of life. Thus, in or igin of life, t r end
wa s : a n a er obi c, ma r i n e pr ok a r yot i c ch emoh et er ot r oph i c l i fe ®a n a e r obi c,
chemoaut r ot r ophs ® aer obic oxygenic phot oaut ot r ophs (cyanobact er ia) ® eukar yot es
® plant s ® animals.
• Nucleic acids having power of self duplicat ion wer e t he fir st biochemical compounds
appear ed in chemical evolut ion. It mar ket s t he beginning of life. Lat er on nucleopr ot ein
and t hen lar ge lipopr ot ein colloidal par t icles called coa cer va t es wer e evolved in
br ot h (hot dilut e soup) of ocean. A coa ce r va t e is a hypot het ical t er m used by Opar in
& Haldane consist ing of nucleopr ot eins, lipids and polysacchar ides which gr ew by
absor bing molecules fr om out side and like bact er ia can divide by budding.
• Coa c e r va t i on: Due t o zwit t er ionic na t ur e, pr ot ein molecules for med colloida l
hydr ophilic complex which got sur r ounded by wat er molecules. These bodies may
separ at e fr om t he body of t he liquid in which t hese ar e suspended (aqueous phase)
and for m a t ype of emulsion. Coalescence of such st r uct ur e pr oduces a separ at ion of
colloids fr om t heir aqueous phase (concer vat ion). These colloid r ich coacer vat es must
been able t o exchange subst ance wit h t heir envir onment and select ively concent r at e
compounds wit hin t hem.
• Fox (1957) synt hesized in vit ro t he pr ot einoids (micr ospher es) by pr olonged heat ing
of 18-20 t ypes of amino acids.
• Th e or y of s p e ci a l cr e a t i on st at es t hat life was cr eat ed by super nat ur al power in
t hat for m which has not under gone any change. It was given by F a t h er Sa u r ez. God
has cr eat ed life in six days fr om media pr ima and man was cr eat ed by him on t he
sixt h day.
• Th e or y of ca t a s t r op h i s m (cat aclysm) was given by Cu vier , accor ding t o which
aft er a gap of cer t ain per iod (called age), t he wor ld under goes a cat ast r ophe (sudden
252 CSIR-NET Life Sciences
calamit y) t o kill almost all t he living or ganisms and t hen God cr eat ed new gener at ion
or new life fr om inor ganic mat t er .
• Th e or y of s p on t a n e ou s ge n e r a t i on (abiogenesis or aut ogenesis) st at es t hat living
beings ar e for med fr om r ain, mud, air , dung and ot her decaying or ganic mat t er . Va n
He lmon t claimed t o pr oduce mice fr om human sweat .
• Th e or y of b i oge n e si s (i.e. life fr om life, omnis vivum ex. vivo) was pr oved by Re d i ,
Sp a l l a n z a n i a nd P a s t e u r independent ly. They dispr oved (r efut ed) t heor y of
spont aneous gener at ion (abiogenesis). Fr ancesco Redi (1668) pr oved t hat flies could
not ar ise fr om put r efying meat wit hout t heir eggs. Spallanzani (1767) demonst r at ed
t hat put r efact ion of meat is due t o micr obes in t he air and it can be pr event ed by
boiling and sealing t he meat in air t ight cont ainer s. Past eur gave a definit e pr oof of
life ar ising fr om pr e-exist ing life using micr obes and st er ilizat ion met hods.
A. Naturalistic theory or chemosynthetic origin (Oparin-Haldane)
Life or iginat ed fr om inor ganic subst ance t hr ough chemical pr ocesses, i .e ., ch e mi ca l e volu t i on
p r eceed s bi ologi ca l evolu t i on . In t he 1920’s, A.I . Op a r i n a n d J .B.S. Ha l d a n e independent ly
post ulat ed t hat t he r educing at mospher e and gr eat er UV r adiat ion on pr imit ive Ear t h favor ed
r eact ions t hat built complex or ganic molecules fr om simple monomer s as building blocks.
Accor ding t o t his t heor y t he fir st living or ganisms ar e pr oduct s of a chemical evolut ion t hat
occur r ed in four st ages:
1. Abiot ic synt hesis and accumulat ion of monomer s, or small or ganic molecules, t hat
ar e t he building blocks for mor e complex molecules
2. J oining of monomer s int o polymer s (e.g., pr ot eins and nucleic acids)
3. For mat ion of pr ot obiont s, named coacer vat es, a colloidal suspension (dr oplet s) which
for med fr om a ggr ega t es of a biot ica lly pr oduced molecules a nd la t er on differ ed
chemically fr om t heir sur r oundings due t o enclosur e of membr ane like st r uct ur e and
named micr ospher es by Sydney fox.
4. Or igin of her edit y dur ing or befor e pr ot obiont appear ance.
Evolutionary Biology 253
The or igin of life was possible in Ear t h’s ancient envir onment :
• Ther e was lit t le at mospher ic oxygen.
• Light ning, volcanic act ivit y, met eor it e bombar dment , and ult r aviolet r adiat ion
wer e mor e int ense.
This is not possible today because
1. Oxygen in Ear t h’s oxidizing envir onment at t acks chemical bonds, r emoving elect r ons.
An impor t ant char act er ist ic of t he ear ly at mospher e must have been t he r ar it y of
oxygen.
2. The moder n at mospher e has a layer of ozone t hat scr eens UV r adiat ion, so t he ener gy
r equir ed t o abiot ically synt hesize or ganic molecules is not available. On pr imit ive
Ear t h, ener gy was available fr om fr equent light ning and int ense UV r adiat ion t hat
penet r at ed t he at mospher e.
Stanley Miller and Harold Urey tested the Oparin/Haldane hypothesis by Experiment
St anley Miller , a gr aduat e st udent in biochemist r y, built t he appar at us shown her e. He filled it
with water (H
2
O) met hane (CH
4
) ammonia (NH
3
) and hydr ogen (H
2
) but no oxygen. He
hypot hesized t hat t his mixt ur e r esembled t he at mospher e of t he ear ly ear t h. (Some ar e not so
sur e). The mixt ur e was kept cir culat ing by cont inuously boiling and t hen condensing t he wat er .
The gases passed t hr ough a chamber cont aining t wo elect r odes wit h a spar k passing bet ween
t hem. At t he end of a week, Miller used paper chr omat ogr aphy t o show t hat t he flask now
cont ained sever al amino acids as well as some ot her or ganic molecules. In t he year s since
Miller ’s wor k, many var iant s of his pr ocedur e have been t r ied. Vir t ually all t he small molecules
t hat ar e associat ed wit h life have been for med: 17 of t he 20 amino acids used in pr ot ein synt hesis,
and all t he pur ines and pyr imidines used in nucleic acid synt hesis. But abiot ic synt hesis of
r ibose — and t hus of nucleosides — has been much mor e difficult .
One difficult y wit h t he pr imeval soup t heor y is how polymer s — t he basis of life it self —
could be assembled. In solut ion, hydr olysis of a gr owing polymer would soon limit t he size it
could r each. Abiot ic synt hesis pr oduces a mixt ur e of L and D enant iomer s. Each inhibit s t he
polymer izat ion of t he ot her . So, for example, t he pr esence of D amino acids inhibit s t he
polymer izat ion of L amino acids (t he ones t hat make up pr ot eins her e on ear t h). This has led t o
a t heor y t hat ear ly polymer s wer e assembled on solid, miner al sur faces t hat pr ot ect ed t hem
fr om degr adat ion, and in t he labor at or y polynucleot ides and polypept ides cont aining about ~50
unit s have been synt hesized on miner al (e.g., clay) sur faces.
Controversy over Classical Oparin-Haldane Theory
The classical t heor y held t hat life or iginat ed as a r esult of such act ions as light ning st r ikes,
put t ing ener gy int o a st r ongly r educing at mospher e, which would r esult in t he for mat ion of
many of t he ‘building blocks of life’ – as shown by t he labor at or y t est s done by Miller . This
t heor y in it s simplest for m has r un int o major pr oblems, and has been pr act ically abandoned by
scient ist s mainly because of t hr ee lines of evidence.
1. Th e a t mos p h e r e wa s p r ob a b ly n ot b y fa r a s r e d u ci n g a s h a d b e e n t h ou gh t (no
fr ee H
2
, NH
3
, CH
4
), but mor e neut r al. The r eact ions t o or ganic mat er ial, fir ed by elect r ic
dischar ge, will st ill t ake place in such an at mospher e, but at a much slower r at e t han in a mor e
r educing at mospher e. At such slow r at es it is ver y difficult t o build up a r at her lar ge r eser voir
of or ganic building blocks, t hus chances of get t ing enough bit s and pieces t o r eact t o big molecules
ar e ver y small: in a ver y dilut e soup – t ype ocean t he molecules never meet in lar ge number s.
254 CSIR-NET Life Sciences
2. Th e t i me a va i la b le for t h e d e ve lop me n t of t h e fi r s t li vi n g ce lls h a s b e e n ge t t i n g
sh or t er a nd shor t er , much less t ha n t he billions of yea r s envisa ged. The big met eor it e
bombar dment t hat also hit t he moon st opped only by about 4.0 Ga. Life developed ear lier would
pr bably have been oblit er at ed by t he impact s. Or ganic mat er ial t hat car r ies t he car bon isot ope
signat ur e of having or iginat ed by phot osynt hesis has been dat ed at 3.85 Ga. The r emains of not
only cell-like or ganisms, but of st r omat olit es, ar e dat ed at 3.5 Ga. St r omat olit es ar e limest ones
secr et ed by t he act ions of phot osynt hesizing bact er ia: not just ver y simple semi-life for ms wer e
ar ound, but fair ly complex bact er ia t hat could per for m t he difficult r eact ion of phot osynt hesis,
i.e., Eubact er ia. The secr et ion of limest one (CaCO
3
) was pr obably mediat ed by t he chemist r y
coupling of r eact ions 1 and 2
H
2
O + CO
2
® CH
2
O + O
2
(React ion 1)
2 HCO
3

+ Ca
2+
® CaCO
3
+ CO
2
+ H
2
O (React ion 2)
If or ganisms use up CO
2
in phot osynt hesis (r eact ion 1), t hey dr ive at t he same t ime
r eact ion 2 t owar ds t he r ight , t hus causing pr ecipit at ion of calcit e.
3. RNA wa s p r ob a b l y t h e fi r s t ge n e t i c ma t e r i a l : An RNA Be gi n n i n g?
Much mor e is known about how incr edibly chemically complex even simple or ganisms
(bact er ia) ar e, and about t he complexit y of inher it ance. All or ganisms alive t oday st or e and
t r ansmit her edit ar y infor mat ion in t wo kinds of molecules called DNA (Deoxyr ibonucleic Acid,
a double spir al shape) and RNA (Ribonucleic Acid, a single spir al shape). Bot h DNA and RNA
ar e made up of four kinds of subunit s called nucleot ides. Sequences of nucleot ides make up t he
genes, and dir ect t he for mat ion of pr ot eins, on which all life depends. Pr ot eins consist of 20
differ ent subunit s called aminoacids, and t he sequence of t he nucleot ides on DNA and RNA
det er mines t he sequence of t he amino-acids in pr ot eins. Ther e ar e lar ge par t s of DNA, however ,
t hat do not code for pr ot eins and appear s t o have no funct ion (‘junk DNA’). Such junk DNA
occur s in Ar chaebact er ia and Eukar yot es, not in Eubact er ia. The for mat ion of t he pr ot eins is
helped along by enzymes, which funct ion as cat alyst s (cat alyst s help a r eact ion along wit hout
par t icipat ing in it ). We call t he nucleot ides and aminoacids t he building blocks of life; bot h have
been found in met eor it es. We have a chicken and egg pr oblem: DNA and RNA “t ell” t he or ganism
how t o make pr ot eins, but t hese same pr ot eins ar e needed t o make DNA and RNA, by act ing as
cat alyst s t o for m t hese big, complex molecules. DNA and RNA ar e ver y complex molecules and
it appear s t o be ver y difficult t o let t hese or iginat e fr om simple amino acids. But t her e is no way
in which we can get pr ot eins t o duplicat e t hemselves.
Mor e and mor e evidence has become available t hat helps in solving t he chicken and egg
pr oblem. It is now known t hat some molecules made of RNA, called r i b ozyme s , can act as
cat alyst s in moder n cells. That means, t hat such RNA molecules could have used bit s and
pieces of t hemselves t o help t hem t o r eplicat e, wit hout any use of pr ot eins. Ther e may t hus
have been a ‘RNA-wor ld’ in which RNA could have per for med t he funct ions of bot h nucleic
acids (DNA, RNA) and pr ot eins. The t heor y t hat t he fir st pr ot o-living t hings wer e RNA-only
or ganisms is becoming widely accept ed (and is called t he t heor y of t he RNA-wor ld).
Sever al ot her bit s of evidence suppor t t his not ion of an or iginal “RNA wor ld”: Many of t he
cofact or s t hat play so many r oles in life ar e based on r ibose; for example: ATP NAD FAD
coenzyme A cyclic AMP GTP. In t he cell, a ll deoxyr ibonucleot ides a r e synt hesized fr om
r ibonucleot ide pr ecur sor s. Many bact er ia cont r ol t he t r anscr ipt ion and/or t r anslat ion of cer t ain
genes wit h RNA molecules (Link t o “r iboswit ches”), not pr ot ein molecules.
Even so, we keep t he pr oblems of a not r educing (neut r al, also not oxidizing) at mospher e,
and ver y lit t le t ime t o get t o t he fir st phot osynt hesizing bact er ia. Ther e ar e t wo main schools
of t hought (wit h many var iant s), oft en combined in one way or anot her .
Evolutionary Biology 255
1. The or iginal sour ce-mat er ial for life was not r eally car bon-based “life”, but some for m
of self-r eplicat ing cr yst al. We ar e looking at : differ ent and innovat ive ways in which
we could have used differ ent commonly occur r ing, nat ur al compounds as some kind
of t emplat e, or even as a r eal par t of ear lier or ganisms. The most commonly cit ed
possible inor ganic compounds ar e pyr it e, FeS
2
(posit ively char ged sur face); and clay
miner als ( negat ively char ged sur face). The float ing bit s and pieces of ‘building blocks
of life’ could have become or ganized in pat t er ns using t he cr yst als as a t emplat e, and
t hey could have st uck fir st t o t he char ged sur face (t he building blocks commonly
have a posit ively and a negat ively char ged end). Not e t hat cr yst als (r egular ly shaped
st r uct ur es) have t he possibilit ies of car r ying “infor mat ion” in t heir ir r egular it ies
(similar t o t he super mar ket bar codes).
2. We looked in t he wr ong place (war m, shallow pond). Reducing cir cumst ances may
not have been aver age in t he at mospher e, but may have occur r ed locally, ar ound
volcanic hot spr ings, and t he fir st for ms of life may have been similar t o chemosynt het ic
bact er ia. Nice ext r a: if life or iginat ed in r at her deep wat er (no sunlight needed): no
pr oblems wit h high r at e of UV-ir r adiat ion in t he absence of an ozone shield (no O
2
).
Addit ionally, many met als t hat ar e r equir ed in ver y small amount s because t hey play
a r ole in complex or ganic molecules (such as chlor ophyll) ar e pr esent in wat er s
st r eaming out of hydr ot her mal vent s – e.g., ir on, nickel, manganese, magnesium,
molybdenum, selenium, copper .
(B). Cosmozoic theory (Theory of panspermia) given by Richter (1865), Helmholtz
(1884), Arrhenius (1908)
They suggest ed t hat life r eached t he ear t h fr om some heavenly body t hr ough met eor it es.
P a n s p e r mi a (pr imit ive for m of life, as suggest ed by Ar r h e n i u s (1908) consist ed of spor es and
micr obes exist ed t hr oughout univer se and pr oduced differ ent for ms of life.
The Murchison Meteorite
This met eor it e, t hat fell near Mur chison, Aust r alia on 28 Sept ember 1969, t ur ned out t o cont ain
a var iet y of or ganic molecules including: pur ines, pyr imidines & polyols — compounds wit h
hydr oxyl gr oups on a backbone of 3 t o 6 car bons such as glycer ol and glycer ic acid. Sugar s ar e
polyols. The amino acids obt ained wer e glycine, alanine, aspar t ic acid, valine and wer e quit e
similar t o t he pr oduct s for med in Miller ’s exper iment s.
The quest ion is: wer e t hese molecules simply t er r est r ial cont aminant s t hat got int o t he
met eor it e aft er it fell t o ear t h. P r ob a b l y n ot : Some of t he samples wer e collect ed on t he same
day it fell and subsequent ly handled wit h gr eat car e t o avoid cont aminat ion. The polyols cont ained
t he isot opes car bon-13 and hydr ogen-2 (deut er ium) in gr eat er amount s t han found her e on
ear t h. The samples lacked cer t ain amino acids t hat ar e found in all ear t hly pr ot eins. Only L
amino acids occur in ear t hly pr ot eins, but t he amino acids in t he met eor it e cont ain bot h D and
L for ms (alt hough L for ms wer e slight ly mor e pr evalent ).
OXYGENATION OF EARTH
Phot osynt hesis (which gener at es fr ee oxygen) or iginat ed in Pr okar yot es (Bact er ia and Ar chaea)
ver y ear ly in Ear t h hist or y: we have evidence fr om st r omat olit es t hat phot osynt hesis occur r ed
about 3.5 Ga ago. The phot osynt hesizing or ganisms wer e pr obably similar t o t he moder n
cyanobact er ia (for mer ly called blue-gr een algae), which ar e Eubact er ia.
256 CSIR-NET Life Sciences
When phot osynt hesizing or ganisms fir st became common, t he fr ee oxygen gas t hat t hey
gener at ed did not immediat ely st ar t t o accumulat e in t he at mospher e: t her e wer e many chemical
compounds ar ound in t he oceans and on land t hat wer e not st able in t he pr esence of fr ee
oxygen gas, and t hat became oxidized.The most common of ‘t hings’ t o be oxidized wer e ir on
(Fe) on land, and sulfur (S) in t he oceans. Pr esent ly, ir on occur s in Fe
2
O
3
(r ust ), sulfur occur s
in sulfat e (SO
4
2-
) in t he oceans. The fr ee oxygen gas was t hus used up in oxidat ion r eact ions for
a consider able t ime, and dur ing t his consider able t ime we t hink t hat t he oxygen concent r at ions
in t he at mospher e did not become higher t han 1 or a few per cent of t he pr esent at mospher ic
level (PAL). Most evidence t hus suggest s t hat t he Ear t h’s at mospher e (t hought not necessar ily
it s oceans) had a level of oxygen equal t o about 10-15% of it s pr esent levels ar ound 2 Ga.
THE GEOLOGIC TIME SCALE AND EVOLUTIONARY RECORD (dates in millions of
years.)
A r emar kable feat ur e of t he t able below is how oft en evolut ionar y changes coincided wit h
geologic changes on t he ear t h. But consider t hat changes in geology (e.g., mount ain for mat ion
or lower ing of t he sea level) cause changes in climat e, and t oget her t hese alt er t he habit at s
available for life. Two t ypes of geologic change seem t o have had especially dr amat ic effect s on
life: cont inent al dr ift and t he impact of ast er oids.
Important Points
• Azoi c Er a –Er a of invisible life.
• Ar ch a e ozoi c Er a -Er a of for mer life or er a of ear ly life.
• P r ot e r ozoi c Er a -Er a of for mer life or Er a of ear ly life.
• P a l a e ozi c Er a -Er a of old life or Er a of ancient life.
• Me s ozoi c Er a -Er a of middle (life also called “age of r ept iles”).
• Coe n ozoi c Er a -Er a of r ecent life (also called “age of mammals, bir ds, insect s and
angiosper ms”).
• Or d ovi ci a n P e r i od : “age of inver t ebr at es”-and or igin of amphibians.
• De von i a n s P e r i od : “age of fishes” and of or igin of amphibians.
• Ca r b on i fe r ou s P e r i od s : “age of amphibians”-and or igin of r ept iles. Ear liest r ept iles
evolved in car bonifer ous per iod.
• Fir st mammals appear ed in Tr i a s s i c p e r i od s .
• J u r a s s i c P e r i od : “age of Rept iles”.
• Holoce n e Ep och : “age of Man”.
Continental Drift
A body of evidence, bot h geological and biological, suppor t s t he conclusion t hat 200 million
year s ago, at t he st ar t of t he Mesozoic er a, all t he cont inent s wer e at t ached t o one anot her in
a single land mass, which has been named Pangaea.
This dr awing of Pangaea (adapt ed fr om dat a of R. S. Diet z and J . C. Holden) is based on a
comput er -gener at ed fit of t he cont inent s as t hey would look if t he sea level wer e lower ed by
6000 feet .
Evolutionary Biology 257
The Geological Time Scale With Dates in Millions of year in bracket
E r a s P e r i od s Ep oc h s Aq u a t i c Li fe Te r r e s t r i a l Li fe
Recent (0.1) Huma ns in t he new
wor ld

Qua t er na r y
(1.8)
Pleist ocene (1.8) Per iodic gla cia t ion
Cont inent a l dr ift
cont inues
Fir st huma ns
Pliocen e Hominids and pongids
Miocene Monkeys a nd a ncest or s
of a pes
Oligocene Ada pt ive r a dia t ion of
bir ds
Eocene


Cenozoic
(65)
Th e Age
of
Ma mma ls

Ter t ia r y
(65)
Pa leocene


All moder n gr oups
pr esent
Ada pt ive r a dia t ion of
ma mma ls

Cr et aceous
(146)
St ill a t t a ched:
N. Amer ica & N.
Eur ope; Aust r a lia
& Ant ar ct ica
Moder n bony fishes,
Ext inct ion of
a mmonit es,
plesiosa ur s,
icht hyosaur s
Ext inct ion of dinosaur s ,
pt er osa ur s Rise of woody
a ngiosper ms, sna kes;
fir st pla cent a l ma mma ls
(Eut her ia )

J ur a ssic
(200)

Afr ica & S.
Amer ica begin t o
dr ift apar t
Plesiosa ur s,
icht hyosa ur s
a bunda nt ; fir st
dia t oms Ammonit es
a ga in a bunda nt
Ska t es, r a ys, a nd
bony fishes
abundant
Dinosa ur s domina nt ;
fir st a ngiosper ms Fir st
ma mma ls;
Ar cha eopt er yx; fir st
liza r ds Ada pt ive
r a dia t ion of dinosa ur s;
insect s abund a nt




Mesozoic
(251)
Th e
Age
Of
Rept iles

Tr ia ssic
(251)
At t he end
Pa nga ea split s
int o La ur a sia a nd
Gondwa na
Fir st plesiosa ur s,
icht hyosa ur s
Ammonit es
a bunda nt a t fir st
Rise of bon y fis h es
Ada pt ive r a dia t ion of
r ept iles:
t hecodont s, t her a psids,
t ur t les,
cr ocodiles, fir st
dinosa ur s

Per mia n (299)
Appa la chia n Mt s.
for med; per iodic
gla cia t ion a nd
a r id clima t e

Ext inct ion of
t r ilobit es,
pla coder ms

Rept iles a bunda nt :
cot ylosa ur s, pelycosa ur s.
Cyca ds, conifer s, ginkgos
Pennsylva nia n
(320)
Ammonit es, bony
fishes
Fir st r ept iles
Coa l swa mps

Ca r bonifer ous

P e r i od i c
a r i d i t y
Mississippia n
(359)
Ada pt ive r a dia t ion of
sha r ks
For est s of lycopsids,
sph en opsids, a nd seed
fer ns
Amphibia ns a bunda nt
La nd sna ils


Pa leozoic
(542)
258 CSIR-NET Life Sciences
Dur ing t he Tr iassic, Pangaea began t o br eak up, fir st int o t wo major land masses: Laur asia
in t he Nor t her n Hemispher e a nd Gondwa na in t he Sout her n Hemispher e. The pr esent
cont inent s separ at ed at int er vals t hr oughout t he r emainder of t he Mesozoic and t hr ough t he
Cenozoic, event ually r eaching t he posit ions t hey have t oday. Let us examine some of t he
evidence.

Devonian
(416)
The "Age of
Fishes"


Ext ensive inland
seas
Placoderms,
car t ilaginous and
bony fishes.
Ammonit es,
naut iloids
Adapt ive radiat ion of
ost r acoder ms,
erypt erids
Ferns, lycopsids, and
sphenopsids
Fir st gymnosperms

Fir st amphibians
Arachnids (scorpions)

Silurian (444)
Mild climat e;
inland seas
Naut iloids, Pilina,
ot her mollusks
Fir st insect s

Ordovician
(488)
Mild climat e,
inland seas
Tr ilobit es abundant
Fir st jawless
vert ebrat es
Fir st fungi
Fir st plant s (liverwort s?)
Fir st millipedes ?

Cambrian
(542)

Per iodic
glaciat ion
Tr ilobit es dominant .
Eurypt erids,
cr ust aceans
mollusks,
echinoderms
sponges, cnidarians,
annelids, and
t unicat es present
No fossils of eukar yot es,
but phylogenet ic t r ees
suggest t hat lichens,
mosses, per haps even
vascular plant s wer e
pr esent .
Pr ot er ozoic
(2500)
Ediacar an
(620)

Fossils rare but
evidence of several
t ypes of
inver t ebr at es
No fossils of
eukaryot es, but
phylogenet ic t r ees
suggest t hat lichens,
mosses, per haps even
vascular plant s wer e
present t owards t he
end.

Evolutionary Biology 259
Shape of the Continents
The east coast of Sout h Amer ica and t he west coast of Afr ica and ar e st r ikingly complement ar y.
This is even mor e dr amat ic when one t r ies t o fit t he cont inent s t oget her using t he boundar ies
of t he cont inent al slopes (e.g., 6000 feet down) r at her t han t he shor elines.
Geology
In bot h miner al cont ent and age, t he r ocks in a r egion on t he east coast of Br azil mat ch
pr ecisely t hose found in Ghana on t he west coast of Afr ica. The low mount ain r anges and r ock
t ypes in New England and east er n Canada appear t o be cont inued in par t s of Gr eat Br it ain,
Fr ance, and Scandinavia. India and t he sout her n par t of Afr ica bot h show evidence of per iodic
glaciat ion dur ing Paleozoic t imes (even t hough bot h ar e now close t o t he equat or ). The pat t er n
of glacial deposit s in t he t wo r egions not only mat ch each ot her but also glacial deposit s found
in Sout h Amer ica, Aust r alia, and Ant ar ct ica.
Fossils
Fossil r ept iles found in Sout h Afr ica ar e also found in Br azil and Ar gent ina. Fossil amphibians
and r ept iles found in Ant ar ct ica ar e also found in Sout h Afr ica, India, and China. Most of t he
mar supial alive t oday ar e confined t o Sout h Amer ica and Aust r alia. But if t hese t wo cont inent s
wer e connect ed by Ant ar ct ica in t he Mesozoic, one might expect t o find fossil mar supials t her e.
In Mar ch 1982, t his pr edict ion was fulfilled wit h t he discover y in Ant ar ct ica of t he r emains of
Polydolops, a 9-ft mar supial.
Rules to Remember
• Wi l l i s t on ’s Ru l e: Dur ing evolut ion of lineage, ser ially homologue par t s t end t o r educe
in number but get mor e and mor e differ ent iat ed e.g. pr awn’s leg.
• Cop e ’s La w: it st at es t hat war m blooded animals t o incr ease in size dur ing t he long
cour se of evolut ion.
• Be r gma n ’s l a w: it st at es t hat war m blooded animals become lar ger in t he nor t her n
and colder par t s of t heir r ange.
• Al l e n ’s l a w: It st at es t hat ext r emit ies (like pinnae, nose) of animals living in t he
nor t her n and colder par t s of t heir r ange have r educed.
• Ga u s e ’s l a w: (Gause, 1934) or t he Comp e t i t i ve Excl u s i on P r i n ci p l e (Har din,
1960).it st at es t hat t wo species having same ecological r equir ement s cannot t o occupy
indefinit ely t he same habit at .
• Gl oge r ’s r u l e: it st at e t hat among war m blooded animals t hose living in war m and
moist climat e develop mor e melanin pigment (ar e dar ker t han animals in cold, dr y
climat es) wher eas for ms in dr y, hot climat es have mor e yellow and r ed pigment .
• J or d a n ’s r u le s : t emper at ur e also influences t he mor phology of cer t ain fishes and is
found t o have some r elat ion wit h t he number of ver t ebr ae. Fishes inhabit ing wat er of
low t emper at ur e t end t o have mor e ver t ebr ae t han t hose of war mer wat er .
260 CSIR-NET Life Sciences
THE HISTORY OF EVOLUTIONARY THEORY
The hist or y of evolut ionar y t heor y is wr it t en in t he life wor ks of many per sons. It cont inues
t oday. A shor t list of ear ly cont r ibuit or s follows.
An a x i ma n d e r (611-547 BC) pr oposed t he t h e o r y o f s p o n t a n e o u s g e n e r a t i o n
(Abiogenesis). It was suppor t ed by Plat o, Ar ist ot le, and Van Helmont .
F a t h er Su a r ez (1548-1617) was t he gr eat est suppor t er of t he t h eor y of sp eci a l cr ea t i on .
F r a n ce s co Re d i (1626-1698) pr oposed t he t h e or y of b i o ge n e s i s . It was supposed by
Spallanzani, Louis Past eur .
Geor ge Lou i s Lecler c (1707-1778) – The Fr ench nat ur alist , also known as t he Cont e de
Buffon, who wr ot e a 44 volume Nat ur al Hist or y of all known plant s and animals. He suggest ed
evolut ion but offer ed no pr oof.
Ca r l Lin n eu s (1707-1778) – A Swedish bot ant ist known as t he “Fat her of Taxonomy”. He
developed t he syst em of binomial nomenclat ur e used t o name and classify or ganisms t oday. He
believed in seper at e cr eat ion and t he fixit y of species. Alt hough he did not belive in evolut ion
his syst emat ic met hods for classifing ar e used t o develop phylogenet ic t r ees.
Geor ge Cu vier (1769-1832) – A ver t ebr at e zoologist . He used compar at ive anat omy t o
classify animals. Pr oposed t he Theor y of Cat ast r ophism which st at ed t hat aft er each wor ldwide
cast ast r ophe t he wor ld was r epopulat ed by t he sur viving species which gave t he appear ance of
cha nce over t ime.
Er a s mu s Da r wi n (1769-1802) – The gr andfat her of Char les Dar win, he was also a
physician and nat ur alist . He suggest ed t he possibilit y of evolut ion based on his st udies of animal
development , ar t ifical br eeding and vest igal or gans.
J a mes Hu t t on (1726- 1797) – Pr oposed t he Unifor mat ion Theor y of Geology. Accor ding
t o Hut t on, t he ear t h is not st at ic but subject t o cont inueous cycles of er osion and uplift ing.
Weat her ed mat er ials wer e deposit ed in layer s which became sediment ar y r ock. This r ock oft en
cont ained fossils and would event ually be lifed fr om t he sea beds t o for m land. It was called t he
unifor mat ion t heor y because t he for ces at wor k wer e t hough t o always act at a unifor m r at e.
We known t hat is not t he case t oday.
Ch a r les Lyell (1796-1875) – Wr ot e t he Pr incipals of Geology which pr ovided suppor t for
Hut t on’s t heor y.
Alfr ed Wegen er 1880-1930) – The Ger man ear t h scient ist who pr oposed t he Theor y of
Cont inent al Dr ift in 1915. Cont r over sial at t he t ime, he pr opose t he cont inent s had under gone
lar ge movement s over t he past 300 million year s. Today t he science of Plat e Tect onics st udies
t he movement of t he ear t h’s cr ust .
Th oma s Ma lt h u s (1766-1834) – Malt hus was an English sociologist who wr ot e an Essay
of Populat ion in 1798. In his essay, he pr oposed t hat deat h and famine wer e inevit able because
t he human populat ion t ended t o incr ease fast er t han t he supply of food. This essay influenced
Dar win gr eat ly and was used in his for mulat ion of t he Theor y of Nat ur al Select ion.
Evolutionary Biology 261
EVIDENCES OF EVOLUTION
1. Fossil Evidence
F os s i l s : Any body, body par t s (t r aces) of animals and veget ables bur ied and pr eser ved by nat ur al
causes. Fossils ar e r emains, t r aces or ot her dir ect evidence of past life for ms. Most fossils for m
fr om bur ial of plant s and animals in sediment ; soft par t s ar e mor e oft en consumed or decomposed
but may leave impr int s if bur ied r apidly. Most fossils ar e embedded in sediment ar y r ock,
wea t her ed pa r t icles t ha t pr ovide st r a t a fr om lower older la yer s t o upper newer la yer s.
Paleont ologist s st udy t he fossil r ecor d based on boundar ies bet ween st r at a, wher e one mix of
fossils gives way t o anot her . Tr ansit ional links ar e int er mediat e bet ween major gr oups. The
fossil r ecor d allows us t o t r ace t he hist or y of t he moder n-day hor se Equus. The ear liest fossils
in t his lineage is Hyr acot her ium , which was t he size of a dog, wit h cusped low-cr owned molar s,
four t oes on each fr ont foot , t hr ee on each hind foot —all adapt at ions for for est living. When
for est s wer e r eplaced by gr asslands, t he int er mediat es wer e select ed for dur able gr inding t eet h,
speed, et c.wit h an incr ease in size and decr ease in t oes.
Types of fossils
(i ) Bod y fos s i l s : har d par t of or ganism such as shell, t oot h, bone et c.e.g. bones of
dinosaur s.
(i i ) Su b fos s i l s : Remains of animals and plant s pr eser ved in r ocks less t han 10,000 year s
e.g. vision in fr ozen ice. Sub fossils wer e for med aft er t he last ice age dur ing Holocene
epoch.
(i i i ) Mi c r ofos s i l s : micr oscopic r emains less t han 0.5 mm or 1/50
t h
inch.
(i v) Ma c r ofos s i l s : Lar ger t han 1 cm in size.
(v) Un u s u a l fos s i l s : Sudden pr eser vat ion of ent ir e or ganism e.g. Solenhofen Limest one
of Sout her n Ger many – cont aining fossils Ar chaeopt er yx.
(vi ) Co p r o l i t i e s : fossils of dr oppings of a nima l fa eca l ma t t er . La r ge copr olit ies of
cr ocodiles, Dinosaur s et c.
(vi i ) Bi oc l a s t : Fossils of fr agment s of fossils enclosed in sediment s. The t er m is usually
applied t o t hin sect ions of fossils under micr oscope.
(vi i i ) Ga s t r ol i t h s : These ar e found in abundance in t he body cavit ies of cer t ain r ept iles.
(i x) P s e u d ofos s i l s : Many object s of inor ganic or igin closely r esemble t he for ms of or ganic
or igin and ar e found in sediment ar y r ock.
Examples
• P r e se r va t i on i n i ce : Woolly mammot hs fr om Siber ia. The flesh is so well pr eser ved
t hat it can be fed t o dogs sever al t housand year s. Discover ed fr om Lena Delt a in 1790
and Siber ia in 1901.
• F os s i l s i n p e t r ol e u m s p r i n gs a n d a s p h a l t s : Rancho La Br ea now in Los Angeles.
• F os s i l s i n r e s i n s a n d a mb e r s : e.g. Fossil fly in amber fr om Balt ic for est s of Eur ope
dur ing Oligocene per iod.
• P t e r i fi ca t i on of h a r d p a r t s : Fossils ar e found in sediment ar y r ocks. Soft par t s
disint egr at e leaving t he fossil por ous. Wat er seeps int o fossil and r eplaces t he har d
262 CSIR-NET Life Sciences
par t par t icle by silica or ir on pyr it e. The pr ocess of so gr adual and slow t hat even t he
out lines of cellular st r uct ur e ar e pr eser ved.
• This pr ocess of fossilizat ion t o pr eser ve t he finer det ails is known as hist omet abasis.
• Mou l d s a n d c a s t s : The mat er ial sur r ounding t he fossil har dens and pr eser ves t he
out er det ails. The act ual bodies disint egr at e and ar e r emoved by slippage of t he gr ound
leaving t he har den cavit ies called moulds. When moulds ar e filled wit h nat ur al deposit s.
They ar e called as cast s e.g. fossils of Pompeii cit y bur ied in volcanic ash of mount
Vesuvious in A.D.79.
• I m p r e s s i o n s: I mpr essions of lea ves of pla nt s, fea t her s of ext inct bir ds, wing
membr anes of flying r ept iles, skin of dinosaur .
• Tr a ck s a n d t r a i l s : The foot pr int s or t r acks left in t he soft moist mud get s har dened
up e.g. t r acks of amphibians discover ed near Pit t sbur g, Ger many fr om Pennsylvanian
per iod.
• Mu mmi e s : Bodies of dead animals or plant s become dehydr at ed in t he deser t s and
ar e pr eser ved as mummies.
La w of Su p e r p os i t i on : t he lower st r at a of a geological for mat ion wer e fir st t o be
deposit ed and is t he oldest .
Con n e ct i n g li n k s ar e t hose living or ganisms which exhibit t he char act er s of t wo differ ent
gr oups of or ganisms. For examples, pr ot opt er us bet ween ost eicht hyes (bony fish) and amphibian;
Or n i t h or -h yn ch u s bet ween r ept ilian and mammalian; Per i p a t u s bet ween annelida and
ar t hr opoda: Fritschiella bet ween aquat ic and land plant s.
P s e u d ofos s i l s – ar e t he fossils like impr essions for med on some r ocks due t o miner al
deposit ion.
Mi s s i n g l i n k s ar e t hose ext inct or ganisms which possessed t he char act er s of t wo differ ent
gr oups of or ganisms. For ex. Archaeopt eryx (lizar d -bir d) is a missing links bet ween t he r ept iles
and t he aves.
Syn a p s i d r e p t i l e s wer e mammals like t hat gave r ise t o mammals. They had a single
t empor al fossa on t he lat er al side of skull and het er odon t eet h. They ar e ext inct .
2. Geological Time Scale
Ge ol ogi ca l t i me s ca l e is t he t abulat e pr esent at ion of sequence and dur at ion of differ ent er as
and per iods wit h t heir dominant for m or life. It had 5 pr inciple er as: P r ot er ozoi c, Ar ch oeozoi c,
P a le ozoi c, Me s ozoi c a n d Ce n ozoi c. Geological hist or y of ear t h is divided int o er as, per iods,
and epochs. Fossil r ecor d pr ovides r elat ive dat ing of r ock layer s; t op layer s of r ock ar e younger
t han lower layer s. Absolut e dat ing met hod uses r adioact ive isot opes. Isot opes each have
par t icular half-life or t ime it t akes for half of isot ope t o decay and become non-r adioact ive.
Car bon-14 (14C) used t o dat e or ganic mat t er ; half decays t o 14N each 5,730 year s; limit ed t o
about last 50,000 year s. Half of pot assium-40 (40K) decays t o ar gon-40 (40Ar ) each 1.3 million
year s; est imat es age of younger r ocks. Ur anium-238 decays t o lead-207; est imat es age of older
r ocks.
3. Mass Extinctions
Ext inct ion is deat h of all member s of species in wild; mass ext inct ions ar e ext inct ions of many
species in shor t t ime. Five mass ext inct ions in fossil r ecor d define end of:
(i ) Or d ovi ci a n
Evolutionary Biology 263
(i i ) Devon i a n
(i i i ) P e r mi a n
(i v) Tr i a s s i c
(v) Cr e t a ce ou s
Following ext inct ions, r emaining gr oups expand t o fill habit at s vacat ed by ext inct species.
Ext i n ct i on of d i n os a u r s a t e n d of Cr e t a ce ou s : Pr oposed in 1977 t hat Cr et aceous ext inct ion
was caused by ast er oid impact . Cr et aceous-Ter t iar y bor der has high level of ir idium, r ar e in
ear t h’s cr ust but common in met eor it es. Calculat ions of effect s of nuclear bomb explosions
(“nuclear wint er ”) compar e wit h wor ldwide climat e cooling expect ed fr om lar ge ast er oid impact .
Wor ldwide layer of soot also defines ir idium layer . Huge met eor it e cr at er of cor r ect age found
in Car ibbean Ocean and Yucat an peninsula; suspect ed sit e of impact of met eor t hat r esult ed in
dinosaur ext inct ion, Mar ine animal fossil r ecor d indicat es mass ext inct ions occur ever y 26
million year s; cor r esponds t o movement of solar syst em wit hin Milky Way galaxy.
The Alvarez Theory
Louis Alvar ez, his son Walt er , and t heir colleagues pr oposed t hat a giant ast er oid or
comet st r iking t he ear t h some 65 million year s ago caused t he massive die-off at t he end of t he
Cr et aceous. Pr esumably, t he impact gener at ed so much dust and gases t hat skies wer e dar kened
all over t he ear t h, phot osynt hesis declined, and wor ldwide t emper at ur es dr opped. The out come
was t hat as many as 75% of all species — including all dinosaur s — became ext inct .
The key piece of evidence for t he Alvar ez hypot hesis was t he finding of t hin deposit s of
clay cont aining t he element ir idium at t he int er face bet ween t he r ocks of t he Cr et aceous and
t hose of t he Ter t iar y per iod (called t he K-T boundar y aft er t he Ger man wor d for Cr et aceous).
Ir idium is a r ar e element on ear t h (alt hough oft en dischar ged fr om volcanoes), but occur s in
cer t ain met eor it es at concent r at ions t housands of t imes gr eat er t han in t he ear t h’s cr ust .
Aft er languishing for many year s, t he Alvar ez t heor y gained st r ong suppor t fr om t he
discover y in t he 90s of t he r emains of a huge (180 km in diamet er ) cr at er in t he Yucat an
Peninsula t hat dat ed t o 65 million year s ago. The abundance of sulfat e-cont aining r ock in t he
region suggests that the im pact generated enorm ous am ounts of sulfur dioxide (SO
2
), which
lat er r et ur ned t o ear t h as a bat h of acid r ain. A smaller cr at er in Iowa, for med at t he same
t ime, many have cont r ibut ed t o t he devast at ion. Per haps dur ing t his per iod t he ear t h passed
t hr ough a swar m of ast er oids or a comet and t he r epeat ed impact s made t he ear t h uninhabit able
for so many cr eat ur es of t he Mesozoic.
4. Biogeographical Evidence
Biogeogr aphy is st udy of dist r ibut ion of plant s and animals t hr oughout t he wor ld.Cur r ent
dist r ibut ion of or ganisms r eflect s evolut ionar y hist or y; or ganisms evolve in one locat ion and
spr ead out int o ot her r egions; for example, no r abbit s ar e found in Sout h Amer ica—t hey
or iginat ed elsewher e and did not each Sout h Amer ica. Physical fact or s, including locat ion of
cont inent s, limit populat ion r ange.
Cont inent al dr ift st at es t hat cont inent s have slowly moved over t ime. Explains close puzzle-
piece fit of east coast of Sout h Amer ica wit h west coast of Afr ica, and ot her cont inent edges,
dist r ibut ion of seed fer ns t hr oughout sout her n cont inent s. Explains dist r ibut ion of ear ly r ept iles
acr oss many cont inent s fr om t ime when land was conjoined and divided dist r ibut ion of mammals
t hat evolved aft er cont inent s par t ed.
264 CSIR-NET Life Sciences
5. Anatomical Evidence
Many or ganisms shar e a unit y of plan; for example, ver t ebr at e for elimbs cont ain same set s of
bones used for differ ent funct ions in bat wings, whale fins, et c. Simplest explanat ion is having
a common ancest or whose basic for elimb plan was modified in succeeding gr oups as each
cont inued along it s own evolut ionar y pat hway.
Homol ogou s or ga n s ar e t hose which ar e similar in or igin and basic st r uct ur e but ar e
adapt ed differ ent t o per for m differ ent funct ions e.g. for e limbs of ver t ebr at es having pent adact yl
limb or igin and similar r esemblance in ar r angement of bones, muscles et c. Ot her examples of
homologous or gans include legs in differ ent insect s’ t eet h of man, t hor n of Bougainvillea and
t endr il of Passiflora.
An a l ogou s or ga n ar e t hose or gans which do per for m similar funct ion but ar e differ ent
in basic st r uct ur e and or igin. For example, wing of insect and wing of bir d leaf of plant and
cladode of Rucus et c.
Ve s t i gi a l s t r u ct u r e s ar e r educed and funct ionless anat omical feat ur es t hat ar e fully
developed and funct ional in ot her ancest r al gr oups. Vest igial st r uct ur es ar e evidence of an
or ganism’s evolut ionar y hist or y. eg. , Flight less bir ds have vest igial wings, Snakes have r emnant s
of a pelvic gir dle, Humans have a t ail bone but no t ail, Whale – femur (lar ge bone). Relat ed
species shar e embr yological development .
(a) All ver t ebr at es exhibit not ochor d dur ing development .
(b) All ver t ebr at es, including humans, exhibit pair ed phar yngeal pouches.
(i) In fishes and amphibians, t hese become funct ioning gills.
(ii) In humans, t hey become t he eust achian t ubes, middle ear cavit y, t onsils, and
t hyr oid and par at hyr oid glands.
(c) Simplest explanat ion is t hat fish not ochor d and phar yngeal pouches ar e pr imit ive
fish feat ur es and fish ar e ancest r al t o ot her ver t ebr at es.
Development of all t r i p l ob l a s t i c animals st ar t s fr om zygot e under goes similar changes
t o for m gast r ula having 3 pr imar y ger m layer s (ect oder m, mesoder m and endoder m) which
have same fat e in or ganogenesis. Ear lier embr yos of differ ent ver t ebr at es r esembles in possessing
similar st r uct ur es like gills slit s, not ochor d, t ail et c. Not t his only in t he cour se of development
at differ ent st ages an embr yo looks like t he embr yo of differ ent phyla for m which t he given
or ganism has evolved. It is t he r eca p i t u la t i on t h eor y (modified int o Biogenet ic law by Haeckel)
which st at es t hat “on t oge n y r e p e a t s p h yloge n y”.
6. Biochemical Evidence
Almost all living or ganisms use t he same basic biochemicals: DNA, ATP, many ident ical enzymes,
DNA t r iplet code, 20 amino acids, int r ons, and hyper var iable r egions. (Pr okar yot es i.e. t r ue
bact er ia (Domain Eukar ya), do not have int r ons. This point s t o a long per iod of t ime since all
living t hings shar ed common acest or y. Similar it y of biochemist r y is explained by descent fr om
common a ncest or .
DNA base sequences differ ences in DNA bet ween a number of or ganisms shows less
differ ence t he mor e closely r elat ed t hey ar e; for example, 2.5% differ ence bet ween humans and
chimpa nzees but 42% differ ence bet ween huma ns a nd lemur s. Amino a cid sequences of
cyt ochr ome c show similar it y bet ween human and monkey, dist ance fr om human t o duck and
gr eat er dist ance t o Candida yeast . Dat a ar e under st andable assuming humans and chimpanzees
shar e a mor e r ecent common ancest or t han do humans and lemur s, ducks, or yeast . Biochemical
evidence is gener ally consist ent wit h anat omical similar it y of or ganisms.
Evolutionary Biology 265
THEORIES OF EVOLUTION
1. Lamark’s theory of inheritance of acquired characteristics
J ea n Ba t i st e d e La ma r ck (1809) pr oposed t he t heor y of “inher it ance of acquir ed char act er s”
in his book “P HI LOSOP HI C ZOOLOGI QUE”. He believed t hat changes in t he envir onment al
fact or s or migr at ion r esult in new disuse of cer t ain ot her or gans. This ext r a use and disuse of
or gans induce t he development and degener at ion r espect ively of cer t ain char act er s. These
change char act er s called acquir ed char act er s ar e inher it ed by next gener at ion and gr adually
accumulat e over many gener at ions leading t o t he or igin of new species. La ma r ck suppor t ed
his t heor y by giving t he examples of evolut ion of long for elimbed and long-necked gir affe fr om
t he deer -like ancest or s. But Lamar ckism was also object ed t o by a number of scient ist s. St r ong
evidence against Lamar ckism was Theor y of Cont inuit y of Ger mplasm pr oposed by Au gu s t
We i s ma n n in 1892 A.D.
2. Evolution by Natural Selection
The idea (given voice by Lamar ck) t hat species could change over t ime was not immediat ely
accept able t o many: t he lack of a mechanism hamper ed t he accept ance of t he idea as did it s
implicat ions r egar ding t he biblical views of cr eat ion. Char les Dar win and Alfr ed Wallace bot h
wor ked independent ly of each ot her , t r aveled ext ensively, and event ually developed similar
ideas about t he change in life over t ime as well as a mechanism for t hat change: n a t u r a l
s e l e ct i on . Char les Dar win secur ed an unpaid posit ion as ship’s nat ur alist on t he H.M.S. Beagle.
The voyage would pr ovide Dar win a unique oppor t unit y t o st udy adapt at ion and gat her a gr eat
deal of pr oof he would lat er incor por at e int o t he t heor y of evolut ion. Dar win spent much t ime
ashor e collect ing plant , animal and fossil specimens, as well as making ext ensive geological
obser vat ions. On his r et ur n t o England in 1836, Dar win began t o cat alog his collect ions and
ponder t he seeming “fit ” of or ganisms t o t heir mode of exist ence. He event ually set t led on four
main point s of t he t heor y.
Fig: Five year voyage of H.M.S. Beagle (1831-1836)
1. Ad a p t a t i on : all or ganisms adapt t o t heir envir onment s.
2. Va r i a t i on : all or ganisms ar e var iable in t heir t r ait s.
3. Ove r -r e p r od u ct i on : all or ganisms t end t o r epr oduce beyond t heir envir onment ’s
capacit y t o suppor t t hem (t his is based on t he wor k of Thomas Malt hus, who st udied
266 CSIR-NET Life Sciences
how populat ions of or ganisms t ended t o gr ow geomet r ically unt il t hey encount er ed a
limit on t heir populat ion size).
4. Since not all or ganisms ar e equally well adapt ed t o t heir envir onment , some will
sur vive and r epr oduce bet t er t han ot her s — t his is known as n a t u r a l s e l e ct i on .
Somet imes t his is also r efer r ed t o as “sur vival of t he fit t est ”. In r ealit y t his mer ely
deals wit h t he r epr oduct ive success of t he or ganisms, not solely t heir r elat ive st r engt h
or speed.
The Wallace-Darwin Theory
• Individuals in a populat ion have var iable levels of agilit y, size, abilit y t o obt ain food,
and differ ent successes in r epr oduct ion.
• Left unchecked, populat ions t end t o expand exponent ially, leading t o a scar cit y of
r esour ces.
• In t he st r uggle for exist ence, some individuals ar e mor e successful t han ot her s,
allowing t hem t o sur vive and r epr oduce.
• Those or ganisms best able t o sur vive and r epr oduce will leave mor e offspr ing t han
t hose unsuccessful individuals.
• Over t ime t her e will be her it able changes in phenot ype (and genot ype) of a species,
r esult ing in a t r ansfor mat ion of t he or iginal species int o a new species similar t o, but
dist inct fr om, it s par ent species.
Support to Natural Selection
1. I n d u s t r i a l me l a n i s m: befor e indust r ial r evolut ion t he dull gr ey for ms of pepper ed
mot h– Biston betularia-wer e dominant t he car bonar ia for m (black) was r ar e because
it was suscept ible t o pr edat ion by bir ds. The r eason was t hat it was conspicuously
visible while r est ing on t r ee t r unks.
The indust r ial r evolut ion r esult ed in lar ge scale smoke which got deposit ed on t r ee
t r unks t ur ning t hem black. Now gr ey var iet ies became suscept ible t he black for ms
flour ished. Replacement of coal by oil and elect r icit y, r educed shoot pr oduct ion – t he
fr equency of gr ey mot hs incr eased again.
2. Si ck l e ce l l a n e mi a a n d ma l a r i a : Individuals homozygous for sickle cell anaemia
die a t a n ea r ly a ge. I n het er ozygous individua ls, t he cell cont a ining a bnor ma l
hemoglobin become sickle shaped, it kills malar ial par asit es effect ively so t hat t hese
individuals ar e able t o come wit h malar ia infect ion much bet t er t han nor mal per son.
3. J . Led er ber g a n d E. Led er ber g pr ovided exper iment al evidence for ‘s e l e ct i on ’ in
ba ct er ia . By using r eplica pla t ing t echniques t hey demonst r a t ed t he pr ocess of
‘s e l e ct i on ’ of ant ibiot ic r esist ance st r ains of bact er ia.
Natural Selection and Genetics (Neo-Darwinism)
Neit her Dar win nor Wallace could explain how evolut ion occur r ed: how wer e t hese inher it able
t r ait s (var iat ions) passed on t o t he next gener at ion? (Recall t hat Gr egor Mendel had yet t o
publish his ideas about genet ics). Dur ing t he 20t h cent ur y, genet ics pr ovided t hat answer , and
was linked t o evolut ion in neoDar winism, also known as t he Moder n Synt hesis.
Evolutionary Biology 267
P op u l a t i on s : Wit hout var iat ion (which ar ises fr om mut at ions of DNA molecules t o pr oduce
new alleles) nat ur al select ion would have not hing on which t o act . A populat ion is a gr oup of
individuals living in t he same geogr aphical ar ea and shar ing a common gene pool. The gene
pool is t he sum of all genet ic infor mat ion car r ied by t he member s of a populat ion.
Al l ge n e t i c va r i a t i on i n a p op u l a t i on i s ge n e r a t e d b y mu t a t i on . Mut at ion is any
her it able change in DNA. Mut at ions can be changes of a single nucleot ide base or may involve
changes in chr omosome number . Whet her a mut at ion is good, neut r al, or har mful depends on
how it affect s sur vival and r epr oduct ive success.
Mu t a t i on Ra t e : Gene mut at ions r esult in new alleles, and ar e t he sour ce of var iat ion
wit hin populat ions. Gene mut at ions ar e ult imat ely behind t he ot her mechanisms t hat pr ovide
var iat ion. Due t o DNA r eplicat ion and DNA r epair mechanisms, mut at ion r at es of individual
genes ar e low, but since each or ganism has many genes, and a populat ion has many individuals,
new mut at ions ar ise in populat ions all t he t ime. Thus, mut at ions ar e r elat ively common, and
t he mut at ion r at e is an adequat e sour ce of new alleles. High levels of molecular var iat ion ar e
common in nat ur al populat ions, alt hough many mut at ions (usually r ecessive) ar e hidden. The
mut at ion r at e var ies gr eat ly among species and even among genes of an individual. Mut at ions
ar e caused by er r or s in DNA r eplicat ion, chemicals, or r adiat ion. Lar ge scale effect s of mut at ion
r esult only when mut at ion is combined wit h ot her fact or s t hat r eshuffle t he gene pool. Select ion
act s on individuals, not t heir individual genes. Sexual r epr oduct ion incr eases var iat ion by
r eshuffling t he genet ic infor mat ion fr om par ent s int o new combinat ions in t heir offspr ing.
Mut at ions pr oduce new alleles.
Additional Sources of Variation
1. Genetic Drift
Gene flow moves alleles among populat ions t hr ough int er br eeding as well as by migr at ion of
br eeding individuals. Gene flow incr eases var iat ion wit hin a populat ion by int r oducing new
alleles pr oduced in anot her populat ion. Cont inued gene flow t ends t o decr ease t he diver sit y
among populat ions, causing gene pools t o become similar . Reduct ion or r est r ict ion of gene flow
bet ween populat ions is essent ial for t he development of new species. The fr equency of alleles
can change fr om gener at ion t o gener at ion as a r esult of chance alone in a small gene pool. This
phenomenon is known as ge n e t i c d r i ft .
Random mat ing involves individuals pair ing by chance, not accor ding t o t heir genot ypes
or phenot ypes. Nonr andom mat ing involves individuals inbr eeding and assor t at ive mat ing.
Inbr eeding is mat ing bet ween r elat ives t o a gr eat er ext ent t han by chance; inbr eeding can
occur if disper sal is so low t hat mat es ar e likely t o be r elat ed and does not change allele
fr equencies, but it does decr ease t he pr opor t ion of het er ozygot es and incr ease t he pr opor t ions
of bot h homozygot es at all gene loci. Assor t at ive mat ing occur s when individuals t end t o mat e
wit h t hose t hat have t he same phenot ype. Assor t at ive mat ing divides a populat ion int o t wo
phenot ypic classes wit h r educed gene exchange. Genet ic dr ift is changes in allele fr equencies of
a gene pool due t o chance or r andom event s. This can occur in lar ge or small populat ions.
Genet ic dr ift causes gene pools of t wo isolat ed populat ions t o become dissimilar as some alleles
ar e lost and ot her ar e fixed. Genet ic dr ift occur s when founder s (or colonizer s) est ablish a new
populat ion, or aft er a genet ic bot t leneck and r esult ant int er br eeding. The founder effect is a
case of genet ic dr ift in which r ar e alleles, or combinat ions of alleles, occur in higher fr equency
in a populat ion isolat ed fr om t he gener al populat ion. Founding individuals cont ain a fr act ion of
t he t ot al genet ic diver sit y of or iginal gene pool. The allele car r ied by founder s is det er mined by
268 CSIR-NET Life Sciences
chance alone. Consider t he Pilgr im colonist s in New England. By no means did t hey r epr esent
all t he genet ic var iat ion of t he human species or even genet ic var iat ions among Eur opeans.
2. A bottleneck effect
Dr ast ic shor t -t er m r educt ions of populat ion size caused by nat ur al disast er s, disease, or pr edat or s
may r esult in (by chance) t he sur vivor s r epr esent ing only a small por t ion of t he or iginal gene
pool. Even when t he populat ion incr eases t o it s or iginal size, a por t ion of it s or iginal genet ic
diver sit y r emains lost . This feat ur e, t er med a bot t len eck , is a pr oblem wit h many endanger ed
species. A bot t leneck effect is genet ic dr ift in which a sever e r educt ion in populat ion size r esult s
fr om nat ur al disast er , pr edat ion, or habit at r educt ion. This r esult s in a sever e r educt ion of t he
t ot al genet ic diver sit y of t he or iginal gene pool. The cheet ah bot t leneck causes r elat ive infer t ilit y
because of t he int ense inbr eeding. Similar ly, t he Hawaiian silver swor d has passed r ecent ly
t hr ough it s own bot t leneck. Recent st udies on humans suggest t hat t her e may have been one
or mor e inst ances of sever e genet ic bot t lenecks in our own pr ehist or y. The bot t leneck effect
pr event s most genot ypes fr om par t icipat ing in pr oduct ion of next gener at ion.
3. Migration (Gene Flow)
Int o or out of a populat ion can br eakdown genet ic differ ences bet ween populat ions. Mut at ions
developing in one populat ion may be spr ead t o ot her populat ions by migr at ion. This ser ves,
like mut at ion, t o int r oduce new alleles int o populat ions.
4. Natural Selection
Not all member s of a populat ion necessar ily have an equal chance of sur viving and r epr oducing
(due t o compet it ion for r esour ces and mat es). By vir t ue of small phenot ypic var iat ions, some
individuals ar e bet t er adapt ed t o t heir envir onment t han ar e ot her s. The bet t er adapt ed
individuals ar e mor e “fit ” and t end t o sur vive and r epr oduce, passing on t heir adapt at ions t o
t he next gener at ion in gr eat er fr equency t han t hose adapt at ions of t he less “fit ” member s of
t he populat ion. Fit ness is a measur e of an individuals abilit y t o sur vive and r epr oduce. Those
wit h t he highest fit ness ar e mor e likely t o sur vive and r epr oduce. Thus, t hey make a gr eat er
cont r ibut ion t o t he gene pool, of t he next gener at ion t han do t hose less “fit ”. Nat ur al select ion
is t he pr ocess of differ ent ial sur vival and r epr oduct ion t hat inevit ably leads t o changes in allele
fr equencies over t ime as t hose individuals who ar e t he most “fit ” sur vive and leave mor e
offspr ing. Ther e ar e t hr ee pat t er ns, or t ypes, of nat ur al select ion.
0 10 20 30 40 50
Medium-
sized
individuals
favored
N
u
m
b
e
r

o
f

i
n
d
i
v
i
d
u
a
l
s

w
i
t
h

p
h
e
n
o
t
y
p
e
Peak gets
higher and
narrower
40 50 30 10 20 0
Selection
0 10 20 30 40 50
Large
individuals
favored
N
u
m
b
e
r

o
f

i
n
d
i
v
i
d
u
a
l
s

w
i
t
h

p
h
e
n
o
t
y
p
e
40 50 30 10 20 0
Peak shifts
in one
direction
Selection
Evolutionary Biology 269
1. Stabilizing Selection
St abilizing select ion favor s t he int er mediat e phenot ype out of a r ange of phenot ypes. The
ext r emes in var iat ion ar e select ed against . Select ion wor ks against bot h ext r emes.
2. Directional Selection
Dir ect ional select ion t ends t o favor phenot ypes at one ext r eme of t he r ange of var iat ion.
Insect icide r esist ance is an example. DDT was a widely used insect icide. Aft er a few year s of
ext ensive use, DDT lost it s effect iveness on insect s. Resist ance t o DDT is a genet ic t r ait t hat
t he pr esence of DDT in t he envir onment made int o a favor ed t r ait . Only t hose insect s r esist ant
t o DDT sur vived, leading over t ime t o populat ions lar gely r esist ant t o DDT. Anot her example
is I n d u s t r i a l me l a n i s m in t he pepper ed mot h (Bist on bet ular ia)). Befor e t he Indust r ial
Revolut ion in t he 18t h and ear ly 19t h cent ur ies, only light -color ed mot hs wer e collect ed in
light -color ed woodlands in England.Ther e was a r ar e, dar k for m. Wit h t he pollut ion caused by
t he bur ing of coal, t he light -color ed t r ee t r unks became dar ker due t o soot . The once r ar e dar k-
color ed mot hs became mor e pr evalent , while t he once-common light -color ed mot hs became
incr easingly r ar e. Reason: pr edat ion by bir ds. The color t hat had t he gr eat est cont r ast wit h t he
backgr ound (t r ee t r unk) was at a disadvant age. Cleanup of t he for est dur ing t he 1950s caused
t he allele fr equencies of light and dar k mot hs t o r ever se t o pr e-Indust r ial Revolut ion levels,
dar k mot hs ar e now r ar e, light mot hs ar e now common. The r esist ance of many bact er ial
species t o ant ibiot ics ia anot her example of dir ect ional select ion. Over 200 species show some
degr ee of ant ibiot ic r esist ance, necessit at ing t he development and mor e pr udent use of a new
gener at ion of ant ibiot ic medicines.
3. Disruptive Selection
In some cir cumst ances, individuals at bot h ext r emes of a r ange of phenot ypes ar e favor ed
over t hose in t he middle. This is called disr upt ive select ion. An example: The r esidues (“t ailings”)
of mines oft en cont ain such high concent r at ions of t oxic met als (e.g., copper , lead) t hat most
plant s ar e unable t o gr ow on t hem. However , some har dy species (e.g. cer t ain gr asses) ar e able
t o spr ead fr om t he sur r ounding uncont aminat ed soil ont o such wast e heaps. These plant s
develop r esist ance t o t he t oxic met als while t heir abilit y t o gr ow on uncont aminat ed soil
decr eases. Because gr asses ar e wind pollinat ed, br eeding bet ween t he r esist ant and nonr esist ant
populat ions goes on. But evident ly, disr upt ive select ion is at wor k. Higher deat h r at es of bot h
less r esist ant plant s gr owing on cont aminat ed soil and mor e r esist ant plant s gr owing on
uncont aminat ed soil leads t o incr easing diver gence of t he populat ions int o t wo subpopulat ions
wit h t he ext r eme manifest at ions of t his t r ait . The evolut ionar y significance of disr upt ive select ion
lies in t he possibilit y t hat t he gene pool may become split int o t wo dist inct gene pools. This may
be a way in which new species ar e for med.
0 10 20 30 40 50
Large and small
individuals favored
N
u
m
b
e
r

o
f

i
n
d
i
v
i
d
u
a
l
s

w
i
t
h

p
h
e
n
o
t
y
p
e
40 50 30 10 20 0
Selection
Two peaks
270 CSIR-NET Life Sciences
Mor t a l i t y Se l e ct i on : Cer t ain genot ypes ar e less successful t han ot her s in sur viving
t hr ough t o t he end of t heir r epr oduct ive per iod.The evolut ionar y impact of mor t alit y select ion
can be felt anyt ime fr om t he for mat ion of a new zygot e t o t he end (if t her e is one) of t he
or ganism’s per iod of fer t ilit y. Mor t alit y select ion is simply anot her way of descr ibing Dar win’s
cr it er ia of fit ness: sur vival.
F e cu n d i t y Se le ct i on : Cer t ain phenot ypes (t hus genot ypes) may make a dispr opor t ionat e
cont r ibut ion t o t he gene pool of t he next gener at ion by pr oducing a dispr opor t ionat e number of
young. Such fecundit y select ion is anot her way of descr ibing anot her cr it er ion of fit ness descr ibed
by Dar win: family size. In each of t hese examples of nat ur al select ion cer t ain phenot ypes ar e
bet t er able t han ot her s t o cont r ibut e t heir genes t o t he next gener at ion. Thus, by Dar win’s
st andar ds, t hey ar e mor e fit . The out come is a gr adual change in t he gene fr equencies in t hat
populat ion.
Mi cr oe vol u t i on : Accumulat ion of small changes in gene pool over a r elat ively shor t
per iod of t wo or mor e gener at ions. In classic obser vat ions and exper iment s, dar k color ed mot hs
went fr om being 10% of populat ion t o 80% when soot color ed t r ees and swit ched success of
pr edat or s.
Place and Sequence of Human Evolution
Ther e is evidence t hat almost all of Hominid evolut ion occur r ed in Afr ica and Asia and t hat t he
evolut ion of t he human species t ook place in Afr ica. Sever al species belonging t o t he genus
Homo can be r ecognized fr om t he fossil r ecor d. For example, Homo habilis lived in Afr ica about
t wo million year s ago and was char act er ized by having a lar ger br ain t hat Australopithecus,
using t ools and being bipedal. Anot her species, Homo erect us appear ed about 1.7 million year s
ago, used fir e and is believed t o have migr at ed t o Asia and Eur ope. Fossils of t he so-called ‘J ava
man’ and ‘Peking man’ belong t o Homo erect us. Homo erect us was r eplaced by Homo sapiens.
A p r i mi t i ve for m of H om o s a p i en s , ca l l e d Ne a n d e r t h a l ma n (H om o s a p i en s
neanderthalensis ), was common in Eur ope and Asia. The Neander t hal men r esembled us, t hough
t hey wer e r elat ively shor t and st ocky and mor e power fully built . The Neander t hals made t ools
and used animal hides as clot hing. They built hut -like st r uct ur es for dwelling and bur ied t heir
dead. Ther e is evidence t hat an abr upt t r ansit ion occur r ed all over Eur ope wher eby t he
Neander t hal man was wiped out and gave way t o t he mor e efficient cousin, t he Cr o-Magnon,
about 34000 year s ago. The Cr o-Magnon people left behind ver y elabor at e cave paint ings showing
t he at t ainment of a for m of cult ur e not unlike our own. Aft er t he last glacial per iod (about
10,000 year s ago), moder n Home sapiens began t o spr ead all over t he globe, cult ivat ed plant s,
domest icat ed animals and r eached enor mous populat ion sizes.
Homo sapiens appear ed in Afr ica about 500,000 year s and pr obably r eplaced Homo erect us
t her e. But in Asia Homo erect us appear s t o have sur vived for anot her 250,000 year s when it
was finally r eplaced by Homo sapiens migr at ing fr om Afr ica.
POPULATION GENETICS
A populat ion is a gr oup of pot ent ially int er br eeding or ganisms of t he same species occupying a
cer t ain ar ea. Member s of a populat ion var y fr om one anot her . This var iat ion is t he r aw mat er ial
on which nat ur al select ion oper at es. Ther e ar e sever al t ypes of mut at ions, bot h at t he gene-
level and t he chr omosome-level. Gene mut at ions pr ovide new alleles, making t hese mut at ions
t he ult imat e sour ce of var iat ion. A gene mut at ion is an alt er at ion in t he DNA nucleot ide
sequence, pr oducing an alt er nat e sequence, t er med an allele.
Evolutionary Biology 271
Mut at ions occur at r andom, and can be beneficial, neut r al, or har mful. Some chr omosomal
mut at ions ar e changes in t he number of chr omosomes inher it ed, while ot her s ar e alt er at ions
in ar r angement of alleles on chr omosomes due t o inver sions and t r anslocat ions.
In sexually r epr oducing or ganisms, genet ic r ecombinat ion is t he r eallocat ion of alleles
and chr omosomes. Recombinat ion r esult s fr om cr ossing-over dur ing meiosis, t he r andom
segr egat ion of chr omosomes t o gamet es dur ing meiot ic division, and t he r andom combinat ion
of gamet es dur ing fer t ilizat ion.
The ent ir e genot ype is subject t o nat ur al select ion since new combinat ions of alleles may
have impr ove t he r epr oduct ive success of t he or ganism. For polygenic t r ait s, t he most favor able
combinat ion may occur when t he r ight alleles gr oup by r ecombinat ion. Not only ar e var iat ions
cr eat ed, t hey ar e also pr eser ved and passed on fr om one gener at ion t o t he next . The gene pool
is t he t ot al of all t he alleles in a populat ion, in t he cont ext of gene fr equencies. Neit her dominance
nor sexual r epr oduct ion will change allele fr equencies.
Har dy-Weinber g equilibr ium is a const ancy of gene pool fr equencies t hat r emains acr oss
gener at ions, and might best be found among st able populat ions wit h no nat ur al select ion or
wher e select ion is st abilizing. Micr oevolut ion is t he accumulat ion of small changes in a gene
pool over a r elat ively shor t per iod.
The Hardy-weinberg Equilibrium
The Har dy-Weinber g equilibr ium is t he cent r al t heor et ical model in populat ion genet ics. The
concept of equilibr ium in t he Har dy-Weinber g model is subject t o t he following hypot heses/
condit ions:
1. The populat ion is panmict ic [couples for m r andomly (panmixia), and t heir gamet es
encount er each ot her r andomly (pangamy)].
2. The populat ion is “infinit e” (ver y lar ge: t o minimize differ ences due t o sampling).
3. Ther e must be no select ion, mut at ion, migr at ion (no allele loss /gain).
4. Successive gener at ions ar e discr et e (no cr osses bet ween differ ent gener at ions).
Under t hese cir cumst ances, t he ge n e t i c d i ve r s i t y of t he populat ion is maint ained and
must t end t owar ds a s t a b l e e q u i l i b r i u m of t he dist r ibut ion of t he genot ype.
The Har dy-Weinber g law can be used under some cir cumst ances t o calculat e genot ype
fr equencies fr om allele fr equencies. Let A1 and A2 be t wo alleles at t he same locus,
p is t he fr equency of allele A1 0 = < p = < 1
q is t he fr equency of allele A2 0 = < q = < 1 and p + q = 1
wher e t he dist r ibut ion of allele fr equencies is t he same in men and women, t hen
p × p = p
2
p × q = pq q × p = pq q × q= q
2
t hey pr ocr eat e : (p + q)
2
= p
2
+ 2pq + q
2
= 1
wher e
p
2
= fr equency of t he A1 A1 genot ype ¬ HOMOZYGOTE
2pq = fr equency of t he A1 A2 genot ype ¬ HETEROZYGOTE
q
2
= fr equency of t he A2 A2 genot ype ¬ HOMOZYGOTE
t hese fr equencies r emain const ant in successive gener at ions.
272 CSIR-NET Life Sciences
Exa mp l e : aut osomal r ecessive inher it ance wit h alleles A and a, and allele fr equencies p
and q:
® fr equency of t he genot ypes: : AA = p
2
and t he phenot ypes [ ]: [A] = p
2
+ 2pq
Aa = 2pq [a] = q
2
aa = q
2
Exa mp l e : phenylket onur ia (r ecessive aut osomal), of which t he delet er ious gene has a
fr equency of 1/100:
® q = 1/100
t her efor e, t he fr equency of t his disease is q
2
= 1/10 000,
and t he fr equency of het er ozygot es is 2pq = 2 × 99/100 × 1/100 = 2/100;
Not e t hat t her e ar e a lot of het er ozygot es: 1/50, t wo hundr ed t imes mor e t han t her e ar e
individuals suffer ing fr om t he condit ion. .
For a r ar e disease, p is ver y lit t le differ ent fr om 1, and t he fr equency of t he het er ozygot es
= 2q.
We use t hese equat ions implicit ly, in for mal genet ics and in t he genet ics of pooled
populat ions, usually wit hout consider ing whet her , and under what condit ions, t hey ar e applicable.
Species Concept and Mechanisms of Speciation
Evolut iona r y t heor y must expla in ma cr oevolut ion, t he or igin of new t a xonomic gr oups
(e.g., new species, new gener a, new families). Speciat ion, or t he or igin of new species, is a
cent r al pr ocess of macr oevolut ion because any genus, family, or higher t axon or iginat es wit h a
new species novel enough t o be t he fir st member of t he higher t axon. The fossil r ecor d pr ovides
evidence for t wo pat t er ns of speciat ion: anagenesis and cladogenesis.
1. An a ge n e s i s (p h yl e t i c e vol u t i on ) = The t r ansfor mat ion of an unbr anched lineage
of or ganisms, somet imes t o a st at e differ ent enough fr om t he ancest r al populat ion t o
just ify r enaming it as a new species.
2. Cl a d oge n e s i s (b r a n ch i n g e vol u t i on ) = The budding of one or mor e new species
fr om a par ent species t hat cont inues t o exist ; is mor e impor t ant t han anagenesis in
life’s hist or y, because it is mor e common and can pr omot e biological diver sit y.
Species ar e oft en consider ed t he br idge bet ween micr o- and macr oevolut ion. Tr adit ionally,
t axonomist s (e.g., Car l Linnaeus) used a t ypological species concept . Individual member s of a
species ar e char act er ized by a par t icular (mor phological) t ype. However , t he pr oblem ar ises
her e t hat t her e is almost always phenot ypic var iat ion wit hin populat ions, which can be ext r eme.
Examples of significant var iat ion caused by single-locus polymor phisms t hat can t hus occur
wit hin one lit t er snow goose and king snake color pat t er ns, var iat ion in dur at ion of cir cadian
cycle in Dr osophila t hr ough var iat ion at t he per locus. The widespr ead exist ence of var iat ion
t oget her wit h Dar win’s view t hat species specific char act er ist ic t r ait s can evolve lead t o an
abandonment of t he t ypological species concept .
The most widespr ead species concept cur r ent ly in use is t he Biological Species Concept
(BSC), a s defined by Er nst Ma yr : Sp e c i e s a r e g r o u p s o f a c t u a l l y o r p o t e n t i a l l y
i n t e r b r e e d i n g p op u la t i on s wh i ch a r e r e p r od u ct i ve ly i s ola t e d fr om ot h e r s u ch gr ou p s .
The cr it er ion her e is t hat int er br eeding or gene exchange among or ganisms must occur ,
eit her act ually or pot ent ially. Viable offspr ing alone is not enough. The t er m pot ent ially r efer s
Evolutionary Biology 273
t o a sit uat ion wher e, for inst ance, t her e ar e populat ions t hat ar e geogr aphically isolat ed but
t hat could int er br eed if br ought in cont act . Such populat ions r epr esent one species under t he
BSC.
Me r i t s of t h e BSC: The definit ion defines species as unit s of evolut ion, by vir t ue of t heir
int er br eeding, which makes it a “nat ur al” species concept fr om an evolut ionar y st andpoint .
P r ob le ms wi t h t h e BSC: It does not apply t o asexual populat ions, t hus r aises t he quest ion
how, for example, many bact er ial species ar e t o be defined.In pr act ice, t he species concept used
for pr okar yot es is st ill a t ypological species concept .
Interbreeding vs. reproductive isolation is not an all-or-none distinction
Exa mp l e s : Hybr id zones of t he hooded cr ow and car r ion cr ow wher e t heir r anges get
int o cont act . These cr ow populat ions ar e neit her fully int er br eeding nor fully isolat ed, hence
t he name semispecies.
Symp a t r i c h yb r i d i za t i on : somet imes species whose r anges have come int o cont act
aft er allopat r ic speciat ion hybr idize t o a limit ed ext ent .
Ri n g s p e ci e s : For inst ance, a califor nian salamander , Ensat ina eschscholt zii, occur s on
bot h sides of t he cent r al valley in Califor nia, but not in t he valley. Ther e is a chain of 7 subspecies
t hat for m a r ing ar ound t he valley. Mit ochondr ial dat a suggest t hat t he or iginat ion of t his
r ange was t he nor t h, fr om which t he populat ions migr at ed sout hwar d at bot h sides of t he
valley. All adjacent populat ions int er br eed, except for t he t wo sout her nmost ones.
P r a ct i ca l l i mi t a t i on s : For allopat r ic populat ions, one would have t o assess whet her t he
member s act ually int er br ed, which is oft en not feasible (t hey might not br eed in capt ivit y).
However , even if possible t o show t hat t hey can int er br eed in capt ivit y, one can not be sur e
t hat t hey would act ually int er br eed in nat ur e when br ought int o cont act .
Ther e ar e many ot her species concept s wit h mor e r est r ict ed usage. One example is t he
p h yloge n e t i c s p e ci e s con ce p t which emphasizes common evolut ionar y hist or y. It is defined
as a ir r educible basal clust er of or ganisms diagnosably differ ent fr om ot her such clust er s, and
wit hin which t her e is a par ent al pat t er n of ancest r y and descent . Any char act er t hat dist inguishes
a populat ion would define it as a species, r egar dless of whet her it int er br eeds wit h ot her s or
not . This species concept will lead t o a classificat ion of or ganisms t hat can somet imes be differ ent
fr om BSC. It also applies t o asexual or ganisms.
Examples for barriers to gene flow
1. Prezygotic barriers
(a) Temporal isolation: Two species of field cr icket s r each r epr oduct ive age in t he fall
and in t he spr ing in t he N East of U.S.
(b) Habitat isolation: Two sympat r ic species of her bivor ous ladybir d beet les feed on t hist les
and blue cohosh, r espect ively. They mat e on t heir r espect ive host plant s, which ar e
t heir micr ohabit at s.
(c) Ethological isolation: Fr equent in animals, t he cour t ing individual sends out signals
only t o conspecific individuals, or , if not , t he cour t ed individual does not r espond.
Female cr ickes t end t o r eact only t o t he conspecific male’s cour t ship song. Male
mot hs ar e oft en only at t r act ed by conspecific’s sex pher omones.
274 CSIR-NET Life Sciences
(d) Mechanical isolation: Two swedish or chids (genus Plat ant her a) use differ ent mot hs
as pollinat or s. Their mor phologies ar e such t hat t he pollen masses (pollinia) at t ach
t o differ ent par t s of t he mot hs, i.e.,t he pr oboscis and t he eye. Plant s wit h int er mediat e
flower mor phology have lower pollinat ion success.
(e) Gametic incompatibility: Especially impor t ant among ext er nal fer t ilizer s, e.g., mar ine
inver t ebr at es. In a species of lar ge mar ine snails, t he sper m cont ains an enzyme t hat
can dissolve t he vit elline membr ane only of a conspecific egg. In flower ing-incompat ible
plant s, when pollen is placed on t he st igma, t he gr owt h of t he pollen t ube t hr ough
t he st yle may be ar r est ed, a phenomenon not well under st ood physiologically.
2. Postzygotic barriers
When pr ezygot ic bar r ier s ar e cr ossed and a hybr id zygot e for ms, one of sever al post zygot ic
bar r ier s may pr event development of a viable, fer t ile hybr id.
(a) Reduced hybrid viability: Genet ic incompat ibilit y bet ween t he t wo species may abor t
development of t he hybr id at some embr yonic st age. For example, sever al species of
fr ogs in t he genus Rana live in t he same r egions and habit at s. They occasionally
hybr idize but t he hybr ids gener ally do not complet e development , and t hose t hat do
ar e fr ail and soon die.
(b) Reduced hybrid fert ilit y: If t wo species mat e and pr oduce hybr id offspr ing t hat ar e
viable, r epr oduct ive isolat ion is int act if t he hybr ids ar e st er ile because genes cannot
flow fr om one species’ gene pool t o t he ot her . One cause of t his bar r ier is t hat if
chr omosomes of t he t wo par ent species differ in number or st r uct ur e, meiosis cannot
pr oduce nor mal gamet es in t he hybr id. The most familiar case is t he mule which is
pr oduced by cr ossing a donkey and a hor se; ver y r ar ely ar e mules able t o backbr eed
wit h eit her par ent species.
(c) Hybrid breakdown: When some species cr oss-mat e, t he fir st gener at ion hybr ids ar e
viable and fer t ile, but when t hese hybr ids mat e wit h one anot her or wit h eit her
par ent species, offspr ing of t he next gener at ion ar e feeble or st er ile. For example,
differ ent cot t on species can pr oduce fer t ile hybr ids, br eakdown occur s in t he next
gener at ion when pr ogeny of t he hybr ids die in t heir seeds or gr ow int o weak defect ive
plant s.
Modes of speciation
Repr oduct ive bar r ier s for m boundar ies ar ound species, and t he evolut ion of t hese bar r ier s is
t he key biological event in t he or igin of new species. An essent ial episode in t he or igin of a
species occur s when t he gene pool of a populat ion is separ at ed fr om ot her populat ions of t he
par ent species. This genet ically isolat ed splint er gr oup can t hen follow it s own evolut ionar y
cour se, as changes in allele fr equencies caused by select ion, genet ic dr ift , and mut at ions occur
undilut ed by gene flow fr om ot her populat ions.
The main modes her e ar e allopat r ic, sympat r ic, par apat r ic, and per ipat r ic speciat ion.
1. Allopatric Speciation
Speciat ion t hat occur s when t he init ial block t o gene flow is a geogr aphical bar r ier t hat physically
isolat es t he populat ion. Such occur r ences include emer gence of mount ain r anges, movement
of glacier s, for mat ion of land br idges, subsidence of lar ge lakes. Geogr aphical populat ions oft en
differ genet ically, which includes genet ic mar ker s associat ed wit h r epr oduct ive isolat ion.
Evolutionary Biology 275
Ad a p t i ve r a d i a t i on = The evolut ion of many diver sely adapt ed species fr om a common
ancest or . Examples of adapt ive r adiat ion ar e t he endemic species (= species confined t o a specific
small geogr aphical r egion) of t he Galapagos Islands which descended fr om small populat ions
which float ed, flew, or wer e blown fr om Sout h Amer ica t o t he islands. Dar win’s finches can be
used t o illust r at e a model for such adapt ive r adiat ion on island chains. A single disper sal event
may have seeded one island wit h a per ipher al isolat e of t he ancest r al finch which diver ged as it
under went allopat r ic speciat ion. A few individuals of t his new species may have r eached
neighbor ing islands, for ming new peipher al isolat es which also speciat ed.
Vi ca r i a n t Sp e ci a t i on : Two populat ions of compar able size ecome separ at ed by a bar r ier .
2. Peripatric Speciation (Founder Effect Speciation, Speciation via “Shifting
Balance”)
Small founder populat ions colonize an ar ea out side t he main populat ion r ange. Populat ions
adapt ed t o an envir onment can be t hought of as occupying peaks in an adapt ive landscape.
These peaks r epr esent alt er nat ive opt imal genot ypic or ganizat ions. The pr oposed mechanism
r elies on a combinat ion of genet ic dr ift (which will be st r ong in small founder populat ions), and
nat ur al select ion. Consider a small kar yot ypic anomaliy in t he par ent al populat ion. If dr ift can
dr ive t his kar yot ype t o a populat ion fr equency > 0.5, it may become t he dominant , and t hus
favor ed kar yot ype. Select ion will t hen dr ive it t o t he near est adapt ive peak. Founder effect
speciat ion is t hought t o cause a r apid r eor ganizat ions of t he genet ic const it ut ion of populat ions,
which is associat ed wit h r epr oduct ive isolat ion. Not ice t he fundament al differ ence bet ween t he
genet ic model for allopat r ic speciat ion in gener al, and for per ipat r ic speciat ion: in per ipat r ic
speciat ion, no envir onment al differ ences in t he t wo habit at s ar e r equir ed.
Evi d e n c e for p e r i p a t r i c s p e c i a t i on : Ma ny exa mples a r e a va ila ble fr om na t ur a l
populat ions, geogr aphic var iat ion of sexual dimor phism (plumage color of male-female) in t he
r obin Pet r oica mult icolor . On mainland Aust r alia, t he dimor phism is unifor m, wher eas on
per ipher al islands, t her e is a wide var iat ion in dimor phism.
Exp e r i me n t a l p op u l a t i on s : Repr oduct ive isolat ion evolves somet imes in populat ions
of Dr osophila and mice t hat ar e subject ed t o r epeat ed populat ion bot t lenecks.
3. Sympatric Speciation
Sympat r ic speciat ion = For mat ion of new species wit hin t he r ange of par ent populat ions.
Repr oduct ive isolat ion evolves wit hout geogr aphical isolat ion. This can occur quickly (in one
gener at ion) if a genet ic change r esult s in a r epr oduct ive bar r ier bet ween t he mut ant s and t he
par ent populat ion. Many plant species have or iginat ed fr om impr oper cell division t hat r esult s
in ext r a set s of chr omosomes-a mut ant condit ion called polyploidy. Depending on t he or igin of
t he ext r a set of chr omosomes, polyploids ar e classified in t wo for ms: aut opolyploids and
allopolyploids.
Aut opolyploid = An or ganism t hat has mor e t han t wo chr omosome set s, all der ived fr om
a single species. For example, Nondisjunct ion in t he ger m cell line (in eit her mit osis or meiosis)
r esult s in diploid gamet es. Self-fer t ilizat ion would double t he chr omosome number t o t he
t et r aploid st at e. Tet r aploids can self-pollinat e or mat e wit h ot her t et r aploids. The mut ant s
cannot int er br eed wit h diploids of t he par ent populat ion because hybr ids would be t r iploid (3n)
and st er ile due t o impair ed meiosis fr om unpair ed chr omosomes. An inst ant aneous special
genet ic event would t hus pr oduce a post zygot ic bar r ier which isolat es t he gene pool of t he
mut ant in just one gener at ion.
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Allopolyploid = A polyploid hybr id r esult ing fr om cont r ibut ions by t wo differ ent species.
Mor e common t han aut opolyploidy. Pot ent ial evolut ion of an allopolyploid begins when t wo
differ ent species int er br eed and a hybr id is pr oduced. Such int er specific hybr ids ar e usually
st er ile, because t he haploid set of chr omosomes fr om one species cannot pair dur ing meiosis
wit h t he haploid set of chr omosomes fr om t he second species. These st er ile hybr ids may act ually
be mor e vigor ous t han t he par ent species and pr opagat e asexually. They would be mor e vigor ous
because t hey have t he best qualit ies of bot h species. At least t wo mechanisms can t r ansfor m
st er ile allopolyploid hybr ids int o fer t ile polyploids. We will discuss just one of t hese:
Dur ing t he hist or y of t he hybr id clone, mit ot ic nondisjunct ion in t he r epr oduct ive t issue
may double t he chr omosome number . The hybr id clone will t hen be able t o pr oduce gamet es
since each chr omosome will have a homologue t o synapse wit h dur ing meiosis. Gamet es fr om
t his fer t ile t et r aploid could unit e and pr oduce a new species of int er br eeding individuals,
r epr oduct ively isolat ed fr om bot h par ent species. Speciat ion of polyploids (especially allopolyploids)
has been ver y impor t ant in plant evolut ion. Some allopolyploids ar e ver y vigor ous because
t hey cont ain t he best qualit ies of bot h par ent species. The accident s r equir ed t o pr oduce t hese
new plant species (int er specific hybr idizat ion coupled wit h nondisjunct ion) have occur r ed oft en
enough t hat bet ween 25% and 50% of all plant species ar e polyploids. Sympat r ic speciat ion may
also occur in animal evolut ion t hr ough differ ent mechanisms.
A gr oup of animals may become isolat ed wit hin t he r ange of a par ent populat ion if genet ic
fact or s cause t hem t o become fixed on r esour ces not used by t he par ent populat ion as a whole.
For example, A par t icular species of wasp pollinat es each species of figs. The wasps mat e and
lay t heir eggs in t he figs. A genet ic change causing wasps t o select a differ ent fig species would
segr egat e mat ing individuals of t he new phenot ype fr om t he par ent al populat ion. Diver gence
could t hen occur aft er such an isolat ion.
4. Parapatric Speciation
Evolut ion of r epr oduct ive isolat ion wit hin one cont inuous populat ion in adjacent r egions. It
ma y r equir e ver y st r ong diver gent select ion wit hin t he popula t ion a nd or select ion for
r epr oduct ive isolat ion it self, which is cont r over sial.
FACTORS FACILITATING SPECIATION
1. Ecological factors
For example, t he low disper sal r at es of Dr osophila and ample oppor t unit ies for geogr aphical
isolat ion may have enabled r apid speciat ion in t he Hawaiian islands.
2. Factors internal to the organism
These ar e in gener al poor ly under st ood, but one inst r uct ive example exist s. Pr imit ive fr ogs
have a shor t inner ear t hat r est r ict s t hem t o a small r ange of fr equencies (and t hus t o a limit ed
pot ent ial t o dist inguish mat ing calls). “Advanced” fr ogs have a much longer inner ear and
fr equency r ange. Speciat ion r at es seem t o have incr eased in fr og evolut ion, which may be
facilit at ed by incr eased oppor t unit ies for sexual select ion, sexual isolat ion, and speciat ion.
3. Adaptive divergence
Two populat ions which adapt t o differ ent envir onment s accumulat e differ ences in t he fr equencies
of alleles and genot ypes. Dur ing t his gr adual adapt ive diver gence of t he t wo gene pools,
Evolutionary Biology 277
r epr oduct ive bar r ier s may evolve bet ween t he t wo populat ions. Evolut ion of r epr oduct ive bar r ier s
would differ ent iat e t hese populat ions int o t wo species. A key point in evolut ion by diver gence is
t ha t r epr oduct ive ba r r ier s ca n a r ise wit hout being fa vor ed dir ect ly by na t ur a l select ion.
Diver gence of t wo populat ions is due t o t heir adapt at ion t o separ at e envir onment s, wit h
r epr oduct ive isolat ion being a secondar y development . Gr adual genet ic diver gence of t wo
populat ions may also r esult in t he evolut ion of pr e-zygot ic bar r ier s. For inst ance, an ecological
bar r ier t o inbr eeding may secondar ily r esult fr om t he adapt at ion of an insect populat ion t o a
new host plant differ ent fr om t he or iginal populat ion’s host .
4. Hybrid zones and the cohesion concept of species
Thr ee possible out comes ar e possible when t wo closely r elat ed populat ions t hat have been
allopat r ic for some t ime come back int o cont act : The t wo populat ions may int er br eed fr eely.
The gene pools would become incor por at ed int o a single pool indicat ing t hat speciat ion had not
occur r ed dur ing t heir t ime of geogr aphical isolat ion. The t wo populat ions may not int er br eed
due t o r epr oduct ive bar r ier s. The gene pools would r emain separ at e due t o t he evolut ionar y
diver gence which occur r ed dur ing t he t ime of geogr aphical isolat ion. Speciat ion has t aken
place. A hybr id zone may be est ablished.
Hybr i d zon e = A r egion wher e t wo r elat ed populat ions t hat diver ged aft er becoming
geogr aphically isolat ed make secondar y cont act and int er br eed wher e t heir geogr aphical r anges
over lap For example, t he r ed-shaft ed flicker of west er n Nor t h Amer ica and t he yellowshaft ed
flicker of cent r al Nor t h Amer ica ar e t wo phenot ypically dist inct populat ions of woodpecker s
t hat int er br eed in a hybr id zone st r et ching fr om sout her n Alaska t o t he Texas panhandle. The
t wo populat ions came int o r enewed cont act a few cent ur ies ago aft er being separ at ed dur ing
t he ice ages. The hybr id zone is r elat ively st able and not expanding.
HOW MUCH GENETIC CHANGE IS REQUIRED FOR SPECIATION?
No gener a liza t ions ca n be ma de a bout genet ic dist a nce bet ween closely r ela t ed species.
Repr oduct ive isolat ion may r esult fr om changes in many loci or only in a few.
Th e p u n ct u a t ed equ i li br i u m mod el has st imulat ed r esear ch on t he t empo of speciat ion.
Tr adit ional evolut ionar y t r ees diagr am t he descent of species fr om ancest r al for ms as br anches
t hat gr adually diver ge wit h each new species evolving cont inuously over long spans of t ime.
Big changes t hus occur due t o t he accumulat ion of many small changes.
Paleont ologist s r ar ely find gr adual t r ansit ions of fossil for ms but oft en obser ve species
appear ing as new for ms suddenly in t he r ock layer s. These species per sist vir t ually unchanged
and t hen disappear as suddenly as t hey appear ed. Even Dar win, who believed species fr om a
common ancest r al st ock evolve differ ences gr adually, was per plexed by t he lack of t r ansit ional
for ms in t he fossil r ecor d. Advocat es of punct uat ed equilibr ium have r edr awn t he evolut ionar y
t r ee t o r epr esent fossil evidence for evolut ion occur r ing in spur t s of r elat ively r apid change
inst ead of gr adual diver gence.
This t heor y was pr oposed by Ni l e s El d r e d ge a n d St e p h e n J ay Gould in 1972. It depict s
species under going most of t heir mor phological modificat ion as t hey fir st separ at e fr om t he
par ent species t hen showing lit t le change as t hey pr oduce addit ional species. In t his t heor y
gr adual change is r eplaced wit h long per iods of st asis punct uat ed wit h episodes of speciat ion.
The or igin of new polyploid plant s t hr ough genome changes is one mechanism of sudden
speciat ion. For a populat ion facing new envir onment al condit ions, genet ic dr ift and nat ur al
278 CSIR-NET Life Sciences
select ion can cause significant change in only a few hundr ed or t housand gener at ions. A few
t housand gener at ions is consider ed r apid in r efer ence t o t he geologic t ime scale.
The fossil r ecor d indicat es t hat successful species sur vive for a few million year s on aver age.
If a species sur vives for five million year s and most of it s mor phological changes occur in t he
fir st 50,000 year s; t hen t he speciat ion episode occur r ed in just 1% of t he species’ lifet ime. Wit h
t his t ime scale, a species will appear suddenly in r ocks of a cer t ain age, linger r elat ively
unchanged for millions of year s, t hen become ext inct .
An evolut ionar y spur t pr eceding a longer per iod of mor phological st asis would explain
why paleont ologist s find so few t r ansit ions in t he fossils r ecor d of a species. Because “sudden”
can r efer t o t housands of year s on t he geological t ime scale, differ ing opinions of punct uat ionalist s
and gr adualist s about t he r at e of speciat ion may be mor e a funct ion of t ime per spect ive t han
concept ual differ ence.
Ther e is clear disagr eement , however , over how much a species changes aft er it s or igin.
Hybr id st er ilit y has a cont inuous r ange, fr om none t o complet e, and may be caused by t wo
main fact or s gener at ion of aneuploid gamet es due t o st r uct ur al chr omosomal differ ences bet ween
t he genes of t he par ent s, which may int er act “dishar moniously”, as opposed t o t he “co-adapt ed”
gene complexes wit hin t he species.
Ar gu me n t s a ga i n s t t h i s h yp ot h e s i s a r e mor phology can change wit hin lineages, st able
lineages in t he fossil r ecor d do fluct uat e, t he low r esolut ion of t he fossil r ecor d (<100000yr ). If
it s r esolut ion was higher , t he ar gument goes, one would obser ve t hat t he appar ent ly sudden
t r ansit ions ar e act ually gr adual t r ansit ions.
P u n ct u a t e d gr a d u a l i s m: Char act er evolut ion is episodic, but not necessar ily associat ed
wit h speciat ion. The most cont ent ious issue her e is bet ween hypot hesis one and t wo. To
dist inguish bet ween t he t wo, one has t o assess whet her mor phological change is usually associat ed
wit h speciat ion.
Mi s s i n g l i n k s : They ar e fr equent in t he fossil r ecor d (e.g., body plan t ypes), and t her e
ar e t wo alt er nat ive possibilit ies for why t hey occur . Fir st , if most major phenot ypic changes
wer e br ought a bout by ma cr omut a t ions (sa lt a t or y evolut ion t hen no links would exist .
Goldschmidt s macr omut at ions wer e pr oposed as a complet e “r epat t er ning” of t he chr omosomal
mat er ial, not single gene mut at ions. Ther e is no evidence for such lar ge scale r epat t er ning
leading t o viable or ganisms. Single gene “macr omut at ions” occur (e.g., homeot ic mut at ions),
but t hey usually ar e highly delet er ious. If most phenot ypic evolut ion occur s accor ding t o t he
punct uat ed equilibr ium hypot hesis via founder effect speciat ion, t he fr equent absence of missing
links would be explicable by t he small t r ansit ional populat ion sizes involved.Thus, t he phenot ypic
changes may be cont inuous, but poor ly pr eser ved.
POLYMORPHISM
A polymor phism is a genet ic var iant t hat appear s in at least 1% of a populat ion. Examples: t he
human ABO blood gr oups, t he human Rh fact or t he human major hist ocompat ibilit y complex
(MHC). By set t ing t he cut off at 1%, it excludes spont aneous mut at ions t hat may have occur r ed
in — and spr ead t hr ough t he descendant s of — a single family.
P r ot e i n P ol ymor p h i s ms : All t he examples above ar e of t he pr ot ein pr oduct s of alleles.
These can be ident ified by: ser ology; t hat is, using ant ibodies t o det ect t he differ ent ver sions of
t he pr ot ein and elect r ophor esis; if amino acid changes in t he pr ot ein alt er it s net elect r ical
char ge, it will migr at e mor e or less r apidly in an elect r ical field.
Evolutionary Biology 279
En zyme s a r e fr e q u e n t l y p ol ymor p h i c: A populat ion may cont ain t wo or mor e var iant s
of an enzyme encoded by a single locus. The var iant s differ slight ly in t heir amino acid sequence
and oft en t his causes t hem t o migr at e differ ent ly under elect r ophor esis. By t r eat ing t he gel
wit h t he subst r at e for t he enzyme, it s pr esence can be visualized. Elect r ophor et ic var iant s of
an enzyme occur r ing in a populat ion ar e called a l l ozyme s .
Re s t r i ct i on F r a gme n t Le n gt h P ol ymor p h i s ms (RF LP s ): Pr ot eins ar e gene pr oduct s
and so polymor phic ver sions ar e simply r eflect ions of allelic differ ences in t he gene; t hat is,
allelic differ ences in DNA. Oft en t hese changes cr eat e new — or abolish old — sit es for r est r ict ion
enzymes t o cut t he DNA. Digest ion wit h t he enzyme t hen pr oduces DNA fr agment s of a differ ent
lengt h. These can be det ect ed by elect r ophor esis.
Si n gl e Nu cl e ot i d e P ol ymor p h i s ms (SNP s ): Development s in DNA sequencing now
make it easy t o look for allelic ver sions of a gene by sequencing samples of t he gene t aken fr om
differ ent member s of a populat ion (or fr om a het er ozygous individual). Alleles whose sequence
r eveals only a single changed nucleot ide ar e called single nucleot ide polymor phisms or SNPs.
SNPs can occur in noncoding par t s of t he gene so t hey would not be seen in t he pr ot ein pr oduct .
might not alt er t he cut t ing sit e for any known r est r ict ion enzymes so t hey would not be seen by
RFLP analysis.
Cop y Nu mb e r P ol ymor p h i s ms (CNP s ): Genet ic analysis (using DNA chips and FISH)
has r evealed anot her class of human polymor phisms. These copy number polymor phisms ar e
lar ge (t housands of base pair s) duplicat ions or delet ions t hat ar e found in some people but not
in ot her s. On aver age, one per son differ s fr om anot her by 11 of t hese. One or mor e have been
found on most chr omosomes, and t he list is pr obably incomplet e. While most of t his DNA is
non-coding, funct ional genes ar e embedded in some of it . How, or if, t he per son adapt s t o t he
r esult ing change in gene number is unknown.
How are polymorphisms useful?
Polymor phism analysis is used: in t i ssu e t ypi n g; 1. in or der t o find t he best mat ch bet ween t he
donor , e.g., of a kidney, and t he r ecipient . 2. finding disease genes. Example: t he gene for
Hunt ingt on’s disease was locat ed when t he pr esence of t he disease was found t o be linked t o a
RFLP whose locat ion on t he chr omosome was known. 3. in populat ion st udies, for example,
assessing t he degr ee of genet ic diver sit y in a populat ion. 4. Det er mining whet her t wo populat ions
r epr esent separ at e species or r aces of t he same species. This is oft en cr it ical t o applying laws
pr ot ect ing endanger ed species. 5. Tr acking migr at ion pat t er ns of a species (e.g., whales).
How do polymorphisms arise and persist?
1. By mu t a t i on . But what keeps them in the population? Sever al fact or s may maint ain
polymor phism in a populat ion.
(a) Founder Effect: If a populat ion began wit h a few individuals — one or mor e of
whom car r ied a par t icular allele — t hat allele may come t o be r epr esent ed in
many of t he descendant s. In t he 1680s, Ar iaant je and Ger r it J ansz emigr at ed
fr om Holland t o Sout h Afr ica, one of t hem br inging along an allele for t he mild
met abolic disease por phyr ia. Today mor e t han 30000 Sout h Afr icans car r y t his
allele and, in ever y case examined, can t r ace it back t o t his couple — a r emar kable
example of t he founder effect .
(b) Genetic Drift: An allele may incr ease — or decr ease — in fr equency simply
t hr ough chance. Not ever y member of t he populat ion will become a par ent and
280 CSIR-NET Life Sciences
not ever y set of par ent s will pr oduce t he same number of offspr ing. The effect ,
called r andom genet ic dr ift , is par t icular ly st r ong in small populat ions (e.g., 100
br eeding pair s or fewer ); when t he gene is neut r al; t hat is, is neit her helpful nor
delet er ious. Event ually t he ent ir e populat ion may become homozygous for t he
allele or — equally likely — t he allele may disappear . Befor e eit her of t hese
fat es occur s, t he allele r epr esent s a polymor phism.
Two examples of reduced polymorphism because of genetic drift
By 1900 hunt ing of t he nor t her n elephant seal off t he Pacific coast had r educed it s populat ion t o
only 20 sur vivor s. Since hunt ing ended, t he populat ion has r ebounded fr om t his populat ion
bot t leneck t o some 100,000 animals t oday. However , t hese animals ar e homozygous at ever y
one of t he gene loci t hat have been examined. Cheet ahs, t he fast est of t he land animals, seem
t o have passed t hr ough a similar per iod of small populat ion size wit h it s accompanying genet ic
dr ift . Examinat ion of 52 differ ent loci has failed t o r eveal any polymor phisms; t hat is, t hese
animals ar e homozygous at all 52 loci. The lack of genet ic var iabilit y is so pr ofound t hat cheet ahs
will accept skin gr aft s fr om each ot her just as ident ical t wins (and inbr ed mouse st r ains) do.
Whet her a populat ion wit h such lit t le genet ic diver sit y can cont inue t o adapt t o a changing
envir onment r emains t o be seen.
Balanced Polymorphism
In r egions of t he wor ld (e.g., par t s of Afr ica) wher e malar ia caused by Plasmodium falciparum
is common, t he allele for sickle-cell hemoglobin is also common. This is because childr en who
inher it one gene for t he “nor mal” bet a chain of hemoglobin and one sickle gene ar e mor e likely
t o sur vive t hat eit her homozygot e. Childr en homozygous for t he sickle allele die young fr om
sickle-cell disease but childr en homozygous for t he “nor mal” bet a chain ar e far mor e suscept ible
t o illness and deat h fr om falcipar um malar ia t han ar e het er ozygot es. Hence t he r elat ively high
fr equency of t he allele in malar ial r egions. When nat ur al select ion favor s het er ozygot es over
bot h homozygot es, t he r esult is b a l a n ce d p ol ymor p h i s m. It account s for t he per sist ence of
an allele even t hough it is delet er ious when homozygous.
An ot h e r e xa mp l e : p r i on p r ot e i n s : All human populat ions ar e polymor phic for t he
pr ion pr ot ein Pr PC. It is encoded by t he pr ion pr ot ein gene (PRNP). Two of t he alleles have
differ ent codons at posit ion 129: one encoding met hionine; t he ot her valine. Homozygosit y for
eit her allele incr eases t he suscept ibilit y t o pr ion diseases. People who ar e het er ozygous ar e
mor e r esist ant . A st udy of elder ly women who had sur vived t he kur u epidemic of t he fir st half
of t he 20t h cent ur y (eat ing t he t issues of t he deceased was banned in 1950) showed t hat 76.7%
of t hem wer e het er ozygot es.
MOLECULAR EVOLUTION
Unt il r ecent ly, evolut ion has been consider ed an or ganismal pr ocess in as much as whole
or ganisms wer e t he unit s of nat ur al select ion and change was evident in populat ions only
insofar as or ganisms in t he populat ion changed. Now it is r eadily appar ent t hat evolut ion can
t ake place at t he molecular level, especially wit hin DNA and RNA molecules, and t hat t his
evolut ion ar ises by pr ocesses ot her t han select ion. For t his r eason, it is somet imes consider ed
non-Dar winian evolut ion.
Evolutionary Biology 281
Evol u t i on a r y ch a n ge d e t e ct a b l e a t t h e mol e cu l a r l e ve l : evolut ion has t r adit ionally
been st udied by compar ing mor phological feat ur es of or ganisms and t heir fossil ancest or s. In
t he 1960’s and 1970’s, it became possible t o look at genet ic change dir ect ly at fir st , gene pr oduct s
(enzymes, nonenzyme pr ot eins, et c.) wer e analyzed using elect r ophor esis or immunological
assay t echniques lat er , DNA and RNA sequences wer e analyzed dir ect ly similar it ies and
differ ences indicat e evolut ionar y r elat ionships.
Ra t e s of Mol e cu l a r Evol u t i on : Ear ly st udies of genet ic var iat ion at t he molecular level
pr oduced sur pr ising r esult s. Fir st , t he genome of most or ganisms is r emar kably diver se, wit h
only a fr act ion of t he genome of eukar yot es consist ing of single-copy genes t hat code for pr ot eins.
Also, genomes ar e not funct ionally int egr at ed; t her e ar e par asit ic sequences and r epet it ive
selfish sequences t hat add not hing t o t he fit ness of t he or ganism wit hin which t hey r eside.
Also, t her e ar e differ ent sequences in differ ent or ganelles of t he cell.
Addit ional st udy of molecular var iat ion indicat ed t hat evolut ionar y change occur s at t his
level just as it does at t he phenot ypic level. However , t he r at es of change at t he t wo levels ar e
fr equent ly differ ent , wit h t he r at e being much higher in cer t ain sit es wit hin genes t han is seen
in or ganisms. How do we account for t his? Ar e t her e differ ent pr ocesses oper at ing at each
level? One of t he most basic t heor ies in evolut ion t o have appear ed since Dar win, called t he
neut r al t heor y of molecular evolut ion, was developed t o account for evolut ion at t he molecular
level. It is also cont r over sial, alt hough not near ly as cont r over sial as Dar win’s t heor y was in
1859.
Neutral Theory of Molecular Evolution
If t her e is a t heor y t hat seems t o be diamet r ically opposed t o Dar winian evolut ion, it is t he
neut r al t heor y of molecular evolut ion. Fir st pr oposed by Kimur a (1968) and elabor at ed fur t her
by King and J ut es (1969) and Kimur a and Oht a (1971), t his t heor y suggest s t hat evolut ionar y
change measur able at t he molecular level can be account ed for in many cases not by select ion,
but by mut at ion and r andom est ablishment of allele fr equencies in populat ions. It does not
oppose t he neo-Dar winian ar gument t hat nat ur al select ion is r esponsible for change at t he
phenot ypic level; t her efor e, mor phological, physiological and behavior al feat ur es of or ganisms
ar e st ill assumed t o ar ise by select ion t hat oper at es on individuals. However , most of t he
var iat ion we obser ve at t he molecular level is assumed t o have lit t le effect on individuals.
The neut r al t heor y suggest s t hat t his ar ises mor e or less const ant ly by mut at ion and
becomes est ablished in populat ions because of r andom r at es of loss and fixat ion. Since evolut ion
at t his level is mor e or less const ant , sequence differ ences bet ween species can ser ve as a
molecular clock which can pr ovide infor mat ion about t he t iming of diver gence of species in a
lineage.
Init ial t est s of t he t heor y made use of allozyme dat a. Br iefly, t his t echnique shows t he
pr esence/absence of var ious allozymes (alt er nat ive elect r ophor et ic for ms of an enzyme coded
for by alt er nat ive alleles) in polymor phic populat ions. Alt hough lit t le is known about what
var iat ion in t he enzymes means in t er ms of funct ion, t hey allow one t o measur e allele fr equencies
in populat ions.
Mor e r ecent dat a fr om DNA sequencing indicat e t hat t he accumulat ion of lar ge amount s
of var iabilit y in noncoding DNA t akes place in many or ganisms (appar ent ly r andomly). Even in
coding r egions (which ar e far less var iable t han noncoding r egions), many molecular changes
ar e appar ent at sit es less impor t ant in det er mining funct ion. This infor mat ion suppor t s t he
neut r al t heor y. However , it is appar ent t hat molecular const r aint s (minimizing changes in
282 CSIR-NET Life Sciences
impor t ant funct ional r egions, for example) ar e influenced st r ongly by select ion. These const r aint s
ar e differ ent for differ ent pr ot eins and help t o explain t he r apid r at es of change of some molecules
and slow r at es of ot her s. Ther efor e, it is not cor r ect t o consider t he Neut r al Theor y and Nat ur al
Select ion t o be alt er nat ive and opposing views in evolut ion. Clear ly, t hey ar e bot h involved and
evidence for bot h is available, no mat t er which point of view (molecular or phenot ypic) is t aken.
The t heor y makes t hr ee basic pr edict ions concer ning molecular evolut ion:
1. r at es of change of genes (and t he pr ot eins t hey code for ) will be const ant over t ime;
2. t he highest r at es of change will be obser ved for t hose por t ions of t he genome t hat ar e
least subject t o funct ional const r aint s, changes in which would lead t o eliminat ion by
select ion; and
3. t he maximum r at e of molecular evolut ion is equal t o t he neut r al mut at ion r at e since
most mut at ions ar e neut r al and any advant ageous mut at ions ar e so r ar e t hey can
have lit t le influence on t he over all r at e of molecular change.
Th e fi r s t p r e d i ct i on (const ant r at es of molecular change) is suppor t ed by molecular
dat a (for example, r at es of change of var ious pr ot eins like hemoglobin and cyt ochr ome c, which
ar e discussed in many t ext s (e.g., Aver s, 1989) t hat has been collect ed since Kimur a fir st began
publishing on t he Neut r al Theor y. The unifor mit y of pr ot ein change wit h t ime (differ ent r at es
for differ ent pr ot eins) suggest s select ion is not a par t icular ly impor t ant fact or because if it
wer e, r at es of change would var y as t he select ion pr essur e changed wit h envir onment al change
(low in const ant envir onment s and high in var iable envir onment s, for example). This also
indicat es t hat t he amount of st r uct ur al change t o pr ot eins pr oduced by var ious t axa can be
used as an indicat or of t he amount of t ime since t he t axa diver ged (molecular clocks). However ,
t he r at e of evolut ion depends on t he DNA sequence examined, and even when t he same sequence
is examined in numer ous lineages, t her e ar e measur able differ ences in r at es. For example,
mit ochondr ial DNA sequences have evolved mor e slowly in t ur t les t han in ot her ver t ebr at es
Th e secon d p r ed i ct i on (r at es of molecular evolut ion will be highest in DNA posit ions
t hat have t he least effect on funct ion) is suppor t ed by most of t he evidence we have fr om
molecular genet ics, for example:
1. int r ons (noncoding r egions of t he genome) var y far mor e t han exons (coding r egions)
in lineages.
2. pseudogenes (nonfunct ional r egions of t he genome cor r esponding t o funct ional r egions)
var y consider ably mor e t han funct ional genes.
3. t he t hir d posit ion of codons var ies mor e t han t he fir st t wo; subst it ut ions at t his t hir d
posit ion ar e called synonymous since t hey do not change t he amino acid coded for by
t he codon.
Th e t h i r d p r e d i ct i on (t he neut r al mut at ion r at e set s t he upper limit t o change; posit ive
select ion for favor able mut at ions is negligible) is suppor t ed by t he obser vat ions given above.
Rat es of sequence change in nonfunct ional r egions of a genome ar e much fast er t han t hose at
funct ionally const r ained r egions. However , t he idea t hat posit ive select ion for favor able changes
cannot influence molecular evolut ion is an ext ension of t he t heor y t hat is not war r ant ed in all
cases. A r ecent t est of t his idea by McDonald & Kr eit man (1991) compar ed sequence dat a fr om
t he alcohol dehydr ogenase gene of t hr ee Dr osophila species, individuals of which wer e sampled
fr om differ ent geogr aphic r egions. They consider ed differ ences obser ved bet ween species as
fixed and differ ences among individuals of t he same species t o be polymor phisms. The neut r al
t heor y pr edict s t hat t he r at io of silent subst it ut ions (which do not alt er t he funct ion of a gene
Evolutionary Biology 283
pr oduct ) t o r eplacement subst it ut ions (which alt er t he amino acid sequence of a pr ot ein) should
be t he same at all sit es even if changes at some sit es ar e funct ionally const r ained; t her efor e,
t he r at io of silent t o r eplacement subst it ut ions will be t he same for bot h polymor phisms (wit hin-
species differ ences) and fixed subst it ut ions (bet ween-species differ ences). What t hey found was
a much higher r at io (29%) of r eplacement t o silent subst it ut ions at fixed sit es t hat dist inguish
species when compar ed wit h polymor phic sit es (5%) which do not dist inguish species. This
suggest s t hat some amino acid-r eplacing subst it ut ions ar e adapt ive and fixed not by dr ift but by
select ion oper at ing on individuals car r ying t hese alt er ed genes. This causes t he fr equency of
t hese alt er ed genes t o incr ease mor e r apidly t han by dr ift alone, and spend less t ime in a
polymor phic st at e t han select ively neut r al silent changes. This cont r adict s one of t he basic
assumpt ions of t he neut r al t heor y.
The neut r al t heor y has engender ed much cont r over sy among populat ion genet icist s, but
has been of immeasur able impor t ance for sever al r easons. Fir st , it account s for much of t he
var iat ion obser ved wit hin and bet ween species at t he molecular level. Second, it fur nishes a
null model for how molecular evolut ion will oper at e in t he absence of select ion
Molecular Clocks
If macr omolecules change const ant ly over t ime, t hen we should expect t hat t he st r uct ur e of
t hese molecules obt ained fr om differ ent or ganisms will be differ ent and t hat t he amount of
differ ences will be an indicat ion of t he lengt h of t ime since t hese or ganisms diver ged fr om a
common ancest or . It is as if a “molecular clock” wer e keeping t ime in or ganisms. This idea has
been used t o est imat e t he t iming of diver gence of var ious gr oups of or ganisms. However , it
depends heavily on t he neut r al t heor y.
One must assume t hat t he molecules in quest ion (or sequences wit hin DNA molecules)
ar e changing r andomly and const ant ly (and t her efor e not in r esponse t o select ion, which would
var y t he r at e). One also needs t o be able t o calibr at e molecular clocks using some r eliable
secondar y sour ce of infor mat ion (just as we use t he news ser vices as a sour ce of “r eal” t ime t o
set our wat ches). This could be fossil evidence indicat ing t he act ual t ime t wo gr oups shar ed a
common ancest or . However , t his is not always available.
These assumpt ions ar e not always met . A basic pr oblem wit h molecular clocks is t hat
t hey do not keep per fect t ime. Ever y gene (or gene pr oduct ) appear s t o change at a r at e
independent of ot her s. For example, cyt ochr ome c and globin genes ar e known t o evolve at
differ ent r at es. Also, differ ent or ganisms exhibit differ ent r at es (molecular evolut ion in humans
is slower t han ot her ant hr opoid apes, for example); and or ganisms in t he same clade may
exhibit differ ent r at es at differ ent t imes.
Also, DNA fr om differ ent genomes evolves at differ ent r at es (mit ochondr ial DNA evolves
at a 10-fold higher r at e t han nuclear DNA; chlor oplast DNA is known t o change ver y slowly;
r DNA has coding r egions t hat ar e highly conser ved and noncoding r egions t hat ar e var iable).
Rat es ar e det er mined dir ect ly by calibr at ion using fossil evidence or indir ect ly by compar ing
wit h an out -gr oup.
Ther e have been many cont r over sies about r at es, however , and t his is one ar ea in which
phylogenies based on differ ent t echniques can differ widely. For example, consider t he concept
of “mit ochondr ial Eve”, a paleolit hic female who lived 200,000 year s go in Afr ica and fr om
whom all pr esent -day humans descended. This is based on a analysis of mit ochondr ial DNA,
which is inher it ed exclusively t hr ough t he female and modified only by mut at ion; it also uses a
st andar d est imat e of mut at ion r at e of 2-4% per million year s. However , t he chr onology and
284 CSIR-NET Life Sciences
or igin ar e disput ed by paleont ologist s, who use fossil and cult ur al evidence t o t r ack human
or igins and consider t he or igin t o be fur t her back in t ime and t o have included mult iple sit es in
Asia, Afr ica and Eur ope.
MOLECULAR ANALYSIS
DNA and DNA pr oduct s (pr ot eins) can be obt ained fr om living plant and animal cells fair ly
easily, and t hen analyzed t o det er mine evolut ionar y r elat ionships among individuals r epr esent ing
var ious t axonomic gr oups. DNA can even be r emoved fr om dr ied plant specimens and fr ozen or
mummified plant and animal t issues, alt hough t her e is usually significant degr adat ion and
t her efor e loss of syst emat ic value.
P r ot e i n e l e ct r op h or e s i s : Because pr ot eins var y in size and have numer ous char ges,
bot h posit ive and negat ive, on t heir sur faces, differ ent pr ot eins will migr at e differ ent ly in a
st ar ch or acr ylamide gel t o which an elect r ic field is applied. Even pr ot eins t hat differ by one
amino acid subst it ut ion will exhibit differ ent posit ions on a gel. These ar e visualized using
subst r at es specific t o t he enzymes of int er est and st ains t o which t he enzymat ic r eact ion binds.
Elect r ophor esis is used t o det er mine t he genot ype of individuals in polymor phic populat ions.
Allozyme elect ophor esis ident ifies allozymes, for ms of an enzyme coded for by differ ent alleles
at a locus. I soz ym e el ect r oph or esi s is similar but isozymes ar e pr oduct s of differ ent loci in a
single genome. This t echnique was mor e widely used befor e DNA isolat ion t echniques wer e
developed, but it is st ill used.
DNA-DNA h yb r i d i za t i on : This met hod basically involves melt ing double-st r anded DNA
of t wo samples at high t emper at ur es (r epr esent at ives of t wo sibling species, for example) and
t hen lower ing t emper at ur es t o r eanneal t he single st r ands fr om each sample (if t hey come
fr om t he same or ganisms, t hey ar e called homoduplexes and if t hey come fr om differ ent
or ganisms, het er oduplexes). Reannealed homoduplexes ar e fair ly t ight ly bound, and r elat ively
high t emper at ur es ar e r equir ed t o melt t hem; however , t he mor e unr elat ed t he t wo wer e
or iginally, melt ing of t he r eannealed het er oduplexes r equir es lower t emper at ur es (if t hey
annealed at all or iginally). This t echnique has been used in many animal st udies (including
human and ot her hominoid gr oups). However , it doesn’t seem t o wor k ver y well wit h plant s
because lit t le r eannealing t akes place.
Re s t r i ct i on F r a gme n t Le n gt h P ol ymor p h i s m (RF LP ): Basically, t his met hod t akes
a molecule of nuclear (or or ganelle) DNA and cut s it at known point s using var ious r est r ict ion
enzymes t hat ar e specific for shor t (usually four or six) sequences of base pair s. The fr agment s
pr oduced by t his cut t ing ar e t hen separ at ed elect r ophor et ically t o pr oduce genet ic mar ker s
t hat can be used t o compar e individuals fr om var ious populat ions. Visualizat ion of t he fr agment s
can be accomplished by eit her r adioact ively labeling t hem or st aining. Dir ect sequencing of t he
fr agment s can also be done if t hey pr ove of syst emat ic value.
Ra n d oml y Amp l i fi e d P ol ymor p h i c DNA (RAP D): This t echnique essent ially uses
shor t synt het ic oligonucleot ide pr imer s t o scan a genome for small inver t ed r epeat s and t hen
amplifies t he sequence in bet ween using PCR (polymer ase chain r eact ion) t o make many copies.
Since t he segment of DNA being amplified is r andomly locat ed and of r andom lengt h (it s sequence
is usually not known), t he sizes (lengt hs) of t hese pieces r esolved on gels fr equent ly var y fr om
genot ype t o genot ype. This var iat ion can be assessed in t he same way as allozyme or RFLP
dat a. RAPD has become especially useful in populat ion genet ics because it can gener at e genet ic
mar ker s t hat ar e polymor phic among individuals in t he same populat ion, so “genet ic individuals”
Evolutionary Biology 285
can be r ecognized in clonal species like fungi (for example, a r ecent RAPD st udy showed t hat
t he wor ld’s lar gest or ganism was a fungus t hat r epr oduced clonally and cover ed many hect ar es
of land). It has also been used t o pr oduce “genet ic finger pr int s” t o r esolve legal disput es. RAPD
is being chosen fr equent ly over ot her molecular t echniques now because it pr oduces r esult s
quickly and r eliably at r elat ively low cost .
Di r e ct Se q u e n ci n g of Ge n ome s : This involves pr ecisely det er mining t he nucleot ide
sequence of a por t ion of a genome. Alt hough t his is mor e labor ious, it pr ovides r elat ively
r eliable dat a wit h which t o est ablish phylogenet ic posit ion of r elat ed t axa. Choices of genome
(or por t ion t her eof) can affect t he r esult s obt ained, but on t he whole sequencing is used (gener ally
in a cladist ic analysis) t o est ablish monophyly or polyphyly of gr oups, ancest r al vs der ived
gr oups, et c.) when phylogenet ic analysis is t he object of t he st udy.
As a gener al r ule, choices of DNA sequences t o be obt ained ar e based on t he amount of
evolut ionar y t ime involved in pr oducing t he var iat ions in t he gr oup under st udy (highly
conser ved por t ions of t he genome ar e used when lar ge amount s of t ime ar e involved and
r apidly-changing por t ions ar e used when not much t ime has passed).
In animals t he mit ochondr ial genome (mt DNA) is fr equent ly sequenced for compar isons
of t axa fr om wit hin t he same or der ; t his is because mt DNA is mat er nally inher it ed, is not
subject t o r ecombinat ion and has numer ous known or t h ogolou s ge n e s (t hat is, homologies of
t he same gene in differ ent t axa; as opposed t o par alogous genes, which ar e differ ent ver sions of
a gene in t he same or ganism).
In plant s, t he chlor oplast genome (cpDNA) is fr equent ly chosen for t he same r easons.
F e w ge n e r a li za t i on s a r e :
1. Molecular t echniques can be applied t o a number of dist inct genomes (nuclear ,
mit ochondr ial, chlor oplast ) using a number of differ ent nucleic acids (r DNA, cpDNA,
var ious RNAs, et c.)
2. Molecular clocks do not keep per fect t ime. Ever y gene (or gene pr oduct ) appear s t o
change at a r at e independent of ot her s. For example, cyt ochr ome c and globin genes
evolve at differ ent r at es. Also, differ ent or ganisms exhibit differ ent r at es for t he
same genes (molecular evolut ion in humans is slower t han ot her ant hr opoid apes, for
example); and or ganisms in t he same clade may exhibit differ ent r at es at differ ent
t imes.
Also, DNA fr om differ ent genomes evolves at differ ent r at es (mit ochondr ial DNA evolves
at a 10-fold higher r at e t han nuclear DNA; chlor oplast DNA is known t o change ver y slowly;
r DNA has coding r egions t hat ar e highly conser ved and noncoding r egions t hat ar e var iable).
Rat es ar e det er mined dir ect ly by calibr at ion using fossil evidence or indir ect ly by compar ing
wit h an out gr oup.
Ther e have been many cont r over sies about r at es, however , and t his is one ar ea in which
phylogenies based on differ ent t echniques can differ widely. For example, consider t he concept
of “mit ochondr ial Eve”, a paleolit hic female who lived 200,000 year s ago in Afr ica and fr om
whom all pr esent -day humans descended. This is based on a analysis of mit ochondr ial DNA,
which is inher it ed exclusively t hr ough t he female and modified only by mut at ion; it also uses a
st andar d est imat e of mut at ion r at e of 2-4% per million year s. However , t he chr onology and
or igin ar e disput ed by paleont ologist s, who use fossil and cult ur al evidence t o t r ack human
or igins and consider t he or igin t o be fur t her back in t ime and t o have included mult iple sit es in
Asia, Afr ica and Eur ope.
286 CSIR-NET Life Sciences
Assumptions
Ra t e of mol e cu l a r ch a n ge i s con s t a n t : Clock can be calibr at ed using a r eliable secondar y
sour ce of infor mat ion (fossils, for example). Assumpt ions not always met (cyt ochr ome c appear s
t o change at differ ent r at es at differ ent t imes
DNA a n a l ys i s : In addit ion t o pr ot eins, DNA is fr equent ly analyzed dir ect ly as a sour ce of
infor mat ion about similar it ies among or ganisms
Molecular Systematics
Use of molecular dat a as char act er s t o assess phylogenet ic posit ion of t axa for classificat ion.
Gener ally uses sequence dat a fr om specific genes (r DNA, mt DNA, cpDNA, et c.). Compar ison of
sequence acr oss t axa r eveals pat t er ns of diver gence in t he past , molecular clocks ar e infer r ed
t o est ablish t iming of diver gence when possible. Choice of genes depends on object ive.
Some DNA changes r elat ively r apidly (mt DNA) and can be used for species wit hin gener a
or gener a wit hin families. Chlor oplast DNA of plant s changes ver y slowly and can be used t o
est ablish phylogenies at t he phylum level as or igin of Angiosper ms, r elat ionships among
gymnosper ms and or igin of land plant s
Ribosomal DNA has bot h highly conser ved r egions and spacer s t hat exhibit r apid change,
so t he choice of r egion depends on t he scale of t he phylogeny t o be r esolved.
Cladist ic analysis is nor mally used t o analyze dat a (which is ver y complex); out gr oups
used t o r oot t r ees
Potential problems
1. molecular clocks var iable
2. same genes must be used for compar ison
Practice Test Paper-I
1. The oldest micr ofossil so far of age 3.5 billion year ago was-
(a) Coa cer va t es (b) Eobiont s
(c) Micr ospher es (d) Cyanobact er ia
2. In his book, “The or igin of life (1938)” opar in submit t ed abiogenesis fir st bur biogenesis
ever since, t his t heor y is named as-
(a) Spont aneous gener at ion (b) Chemical or igin
(c) Pr imar y abiogenesis (d) Biogenesis
3. Exper iment al evidence for molecular evolut ion of life was pr ovided by-
(a) Opar in (b) Haldane
(c) Ur ey and Miller (d) Syndey fox
4. Dur ing pr e-biot ic or igin of life which chemical played impor t ant r ole in for mat ion of
nucleot ide specially guanosine-
(a) CH
4
(b) CO
2
(c) NH
3
(d) HCN
5. Among t he following which molecule t ill now not synt hesized by mimicking t he
envir onment of pr e-biot ic envir onment -
(a) Ribose (b) Pyr imidines
(c) Pur ine (d) L-aminoacids
6. Among t he following t he evidence of evolut ion fr om biogeogr aphy is-
(a) Embr yo development (b) Plat e t ect onics
(c) Dar win finches (d) Dar win t ur t les
7. Thor ns of Bougenwalia plant and t endr il of cucur bit s ar e-
(a) Homologous or gans (b) Par alogous or gan
(c) Analogous or gan (d) Or t hologous or gan
8. Placent al mammals such as mouse, wolf, Aust r alian mar supials such as mar supial
mouse, Tasmanian wolf shows-
(a) Par allel evolut ion (b) Conver gent evolut ion
(c) Diver gent evolut ion (d) Phylet ic evolut ion
9. Which of t he following is not an vest igial or gan in humans-
(a) Ear muscles (b) Tail ver t ebr a
(c) Pr emolar (d) Appendix
10. Which of t he following was ear liest for m wit h lipid bilayer and can r epr oduce by
budding-
(a) Coa cer va t es (b) Micr o spher es
(c) pr ot obiont s (d) Monospher es
288 CSIR-NET Life Sciences
11. Biogenet ic law of Von Baer & Ear nst haekel is
(a) Phylogeny r epeat s ont ogeny
(b) Ont ogeny r epeat s phylogeny
(c) Ont ogene r epeat s phyt ogene
(d) Ont ogeny and phylogeny ar e cyclic
12. Evidence fr om fossils r ecor ds ar e obt ained by calculat ing age of fossil found in–
(a) Met amor phic r ock (b) Sediment ar y r ocks
(c) Igneous r ocks (d) Ear t h cr ust
13. Mammals or iginat ed dur ing t he per iod-
(a) Tr iassic (b) J ur assic
(c) Cr et aceous (d) Per mian
14. Fir st plant having seed habit (Het er ospor ous Pt er odophyt e) or iginat ed dur ing-
(a) Silur ian (b) Devonian
(c) Car bonifer ous (d) Per mian
15. Fir st human appear ed dur ing-
(a) Oligocene (b) Miocene
(c) Pliocene (d) Pleist ocene
16. Er a of r ept iles and gymnosper m is-
(a) Pr ecambr ian (b) Paleozoic
(c) Mesozoic (d) Cenozoic
17. The cor r ect or der of evolut ion of hor se is-
(a) Mesohippus, Hyr acot her ium, Mer yhippus, pliohippus, equus.
(b) Mesohippus, Mer yhippus, Hyr acot her ium, pliohippus, equus.
(c) Mesohippus, Mer yhippus, pliohippus, Hyr acot her ium, equus.
(d) Hyr acot her ium, Mesohippus, Mer yhippus, pliohippus, equus.
18. Dar win’s t heor y of pangenesis was r efut ed by-
(a) Recapit ulat ion t heor y (b) t heor y of Ger mplasm
(c) Chr omosome t heor y (d) t heor y of biogenesis
19. Mut at ion t heor y of Hugo de vr ies was put for war d while wor king on-
(a) Drosophila (b) Ancon sheep
(c) Oenot hera amarckiana (d) Ant irrhinum
20. Evolut ion at genet ic level is t er med as-
(a) Micr oevolut ion (b) Macr oevolut ion
(c) Gene Evolut ion (d) Point mut at ion
21. Which of t hem do not cause var iat ion at genet ic level-
(a) Mut at ion and r ecombinat ion
(b) Gene migr at ion and dr ift
(c) Nat ur al select ion and ar t ificial select ion
(d) Panmict ic populat ion
Practice Test Paper–Evolutionary Biology 289
22. The r aw mat er ial for evolut ion is var iabilit y of gene or allele at /in-
(a) individual level (b) populat ion
(c) gene pool (d) communit y
23. Founder effect is concer ned wit h-
(a) Gene migr at ion (b) Genet ic dr ift
(c) Nat ur al select ion (d) Mut at ion
24. If t he individual ar one ext r eme of t he size dist r ibut ion (eg., lar ger one ) cont r ibut e
mor e offspr ing t o next gener at ion t hen such select ion is called as-
(a) Dir ect ional
(b) Disr upt ive
(c) Cyclic
(d) St abilizing
25. Examples of polymor phism in human is-
(a) ABO blood gr oup (b) Sickle cell anaemia
(c) height and Int elligence (d) All of t he above
26. When t he pr eser vat ion of genet ic var iabilit y is t hr ough het er ozygot e super ior it y it is
t er med as-
(a) Het er opolymor phism (b) Balanced polymor phism
(c) St abilizing polymor phism (d) Dir ect ional polymor phism
27. Type of speciat ion due t o polyploidy is-
(a) Allopat hic (b) Par apat r ic
(c) Per ipat r ic (d) Sympat r ic
28. When t he t wo species ar e mor phologically almost ident ical but r epr oduct ively isolat ed,
ar e t er med as-
(a) Taxonomic species (b) Ecot ypes
(c) Sibling species (d) Mor phospecies
29. Met hanogens ar e found in-
(a) Mar shy ar eas (b) Flor a of cat t le r umen
(c) Biogas fer ment or (d) All such places
30. Each of us is par t of t he ongoing evolut ion of t he human species. Which of t he following
occur r ences would have t he gr eat est impact on t he fut ur e biological evolut ion of t he
human populat ion?
(a) You wor k out ever y day so t hat you st ay physically fit and healt hy.
(b) A mut at ion occur s in one of your skin cells.
(c) You move t o Hawaii, t he st at e wit h t he longest life expect ancy.
(d) A mut at ion occur s in one of your sper m or egg cells.
31. The pr ocesses of ____ and ____ gener at e var iat ion, and____ pr oduces adapt at ion t o
t he envir onment .
(a) sexual r ecombinat ion . . . nat ur al select ion . . . mut at ion
(b) mut at ion . . . sexual r ecombinat ion . . . genet ic dr ift
290 CSIR-NET Life Sciences
(c) genet ic dr ift . . . mut at ion . . . sexual r ecombinat ion
(d) mut at ion . . . sexual r ecombinat ion . . . nat ur al select ion
32. Bir ds wit h aver age-sized wings sur vived a sever e st or m mor e successfully t han ot her
bir ds in t he same populat ion wit h longer or shor t er wings. This illust r at es
(a) t he founder effect (b) st abilizing select ion
(c) ar t ificial select ion (d) gene flow
33. Which of t he following scenar ios would most likely r esult in micr oevolut ion of a
populat ion of humans?
(a) Only r andom mat ing t ook place in all people t hat r epr oduced in Nor t h Amer ica
(b) A colony of humans on t he moon was isolat ed fr om ear t h
(c) The incidence of skin cancer in adult s over age 40 r ose significant ly
(d) Bot h a and c
34. Which of t he following is t he best example of gene flow.
(a) A polyploid plant develops
(b) Genes ar e shuffled by cr ossing over of chr omosomes dur ing meiosis
(c) An ear t hquake r esult s in t he for mat ion of a canyon split t ing a populat ion of
t oads apar t
(d) Wind blows pollen fr om one populat ion of plant s t o anot her and cr oss fer t ilizat ion
occur s
35. St abilizing select ion
(a) favor s int er mediat e var iant s in a populat ion
(b) pr event s mut at ions fr om occur r ing
(c) occur s when some individuals migr at e t o an ar ea wit h differ ent envir onment al
condit ions
(d) can only t ake place in species exhibit ing sexual dimor phism
36. Accor ding t o t he Har dy-Weinber g t heor em, t he fr equencies of alleles in a populat ion
would r emain const ant if ______________ is t he only pr ocess t hat affect s t he gene
pool.
(a) mut at ion (b) genet ic dr ift
(c) sexual r epr oduct ion (d) micr oevolut ion
37. In t he Har dy-Weinber g t heor em p2 r epr esent s
(a) t he t ot al alleles in t he gene pool
(b) t he het er ozygous dominant s in t he gene pool
(c) t he homozygous r ecessives in t he gene pool
(d) t he homozygous dominant s in t he gene pool
38. In t he Har dy-Weinber g t heor em, 1 r epr esent s
(a) The t ot al alleles in t he gene pool
(b) t he het er ozygous dominant s in t he gene pool
(c) t he homozygous r ecessives in t he gene pool
(d) all t he possible phenot ypes in t he gene pool
Practice Test Paper–Evolutionary Biology 291
39. Two animals ar e consider ed of differ ent species if t hey
(a) look differ ent (b) cannot int er br eed
(c) live in differ ent h (d) ar e geogr aphically isolat ed
40. Which of t he following is t he fir st st ep in allopat r ic speciat ion?
(a) genet ic dr ift (b) geogr aphical isolat ion
(c) polyploidy (d) hybr idizat ion
41. Most of t he t ime, species ar e ident ified by t heir appear ance. Why?
(a) If t wo or ganisms look alike, t hey must be t he same species
(b) This is t he cr it er ion used t o define a biological species
(c) If t wo or ganisms look differ ent , t hey must be differ ent species
(d) This is t he most convenient way of ident ifying species
42. A new species can ar ise in a single gener at ion
(a) t hr ough geogr aphical isolat ion
(b) in a ver y lar ge populat ion t hat is spr ead over a lar ge ar ea
(c) if a change in chr omosome number cr eat es a r epr oduct ive bar r ier
(d) if allopat r ic speciat ion occur s
43. The evolut ion of numer ous species, such as Dar win’s finches, fr om a single ancest or
is called
(a) adapt ive r adiat ion (b) sympat r ic speciat ion
(c) gr adualism (d) nondisjunct ion
44. Accor ding t o t he _________ model, evolut ion occur s in spur t s; species evolve r elat ively
r apidly, t hen r emain unchanged for long per iods.
(a) nondisjunct ion (b) gr adualist
(c) adapt ive r adiat ion (d) punct uat ed equilibr ium
45. Sympat r ic speciat ion is
(a) t he appear ance of a new species in t he same ar ea as t he par ent populat ion.
(b) init iat ed by t he appear ance of a geogr aphical bar r ier
(c) t he emer gence of many species fr om a single ancest or
(d) especially impor t ant in t he evolut ion of island species
46. In some animals eg. Axolat al lar vae of Ambyst oma, t he lar vae fails t o under go
met amor phosis (iodine deficiency). It develops gonads, at t ains sexual mat ur it y and
st ar t r epr oduct ion. This is called-
(a) Par t henogenesis (b) Neot eny or paedogenesis
(c) Ret r ogr ess met amor ph (d) St agnant met amor phosis
47. Dar win finches fr om var ious islands of Galapagos island differ ed in size and shape of
bill due t o-
(a) Mut at ion (b) Adapt ive r adiat ion
(c) Compet it ion (d) Gene migr at ion
292 CSIR-NET Life Sciences
48. The enzyme which is occur s in all eukar yot es and show gr eat homology is-
(a) Cyt ochr ome oxidase (b) Cyt ochr ome c
(c) Cyt ochr ome b (d) Cyt ochr ome a
49. The hor mone which is almost ident ical and occur s in all ver t ebr at es is-
(a) Insulin (b) Thyr oxine
(c) Glucagon (d) ACTH
50. The phylogenict ically closed r elat ive of humans ar e-
(a) Langur s (b) Apes
(c) Mokeys (d) Shr ews
51. Which of t he following is not dar win’s post ulat e-
(a) St r uggle for exist ence
(b) sur vival of fit t est
(c) or igin of new species
(d) Inher it ence of Acquir ed char act er
52. Which is main cause of genet ic var iat ion at individual level-
(a) Gene mut at ion (b) Cr ossing over
(c) Fer t ilizat ion & meiosis (d) All if t he above
53. Tot al gene cont ent of individual populat ion is t er med as-
(a) Genome (b) Gene pool
(c) Gene bank (d) Genot ype
54. A mut at ion ar ising in small populat ion is eit her fixed or lost just by chance ir r espect ive
of it s adapt ive value is t er med as-
(a) Silent t heor y (b) Genet ic dr ift
(c) Ar t ificial select ion (d) Dir ect ional select ion
55. The main r eason for evolut ion of new species is-
(a) Geogr aphical isolat ion (b) Nat ur al select ion
(c) Repr oduct ive bar r ier (d) Mut at ion
56. Biological met hod for clean up of cont aminat ed soil and gr ound wat er is t er med as-
(a) Bior est or at ion (b) Biomagnificat ion
(c) Bior emediat ion (d) Biosor pt ion
57. Ar chaebact er ia gr ows at t emper at ur e-
(a) 0-20
o
C (b) 30-40
o
C
(c) 50-120
o
C (d) All such t emper at ur es
58. Agar -agar used as solidifying agent in bact er iological media is obt ained fr om-
(a) Spir ulina (b) Gelidium
(c) Fucus (d) Por phyr a
Practice Test Paper–Evolutionary Biology 293
59. Which of t he following or ganism can car r y out t he r eact ion-
NO
2
+ oxygen ® ®NO
3
+ e n e r gy
(a) Nit rosomanas (b) Pseudomonas rubrum
(c) Micrococcus denit rificans (d) Nit robact er
60. Evolut ion of moder n wheat Trit icum aest ivum is a r esult of-
(a) Mut at ion (b) Aut opolyploidy
(c) Allopolyploidy (d) Segment al polyploidy
61. Pr ot opor phyr in, t he ancient solar ener gy t r apping molecules ar e made fr om-
(a) Succinic acid & glycine (b) Alanine & glut amic acid
(c) Lysine & pr aline (d) Acet yl coA & oxaloacet at e
62. Among t he following which do not br ing var iat ion at Individual level-
(a) Gene mut at ion
(b) Fer t ilizat ion
(c) Meiosis and Cr ossing over
(d) Chr omosome aber r at ion and Hybr idizat ions
63. Nat ur al select ion will oper at e under condit ion-
(a) Mut at ional equilibr ium
(b) Random mat t ing
(c) Differ ent ial r epr oduct ion
(d) Equal chances for all genot ypes t o live and r epr oduce
64. Nat ur al select ion will not oper at e if-
(a) Populat ion is isolat ed and small
(b) Mut at ing populat ion
(c) Random mat t ing populat ion
(d) Lar ge populat ion
65. Dist inct and st able phenot ypes wit hin species ar e t er med-
(a) Polymor phic var iat ion (b) Cr ypt ic var iat ion
(c) Geogr aphical var iat ion (d) Local var iat ion
66. The main cause of blast ogenic and somat ogenic var iat ion-
(a) Endocr ine glands (b) Mut at ion
(c) Nat ur al select ion (d) Panmict ic r epd.
67. Cr ossing of hybr id t o it s par ent s is called-
(a) Par ent al hybr idizat ion (b) Int er ogr essive hybr d.
(c) Back hybr idizat ion (d) St abilizing hybr d.
68. Bot h Tr ansit ion and Tr ansver sion t ype of mut at ion is induced by-
(a) Met hyl met hane sulfat e (b) Nit r ous acid
(c) Base analogues (d) Acr adines
294 CSIR-NET Life Sciences
69. Which is best exper iment al demonst r at ion t o pr ove nat ur al select ion-
(a) Leder ber g’s r eplica r eplicat ing
(b) Ket t elwell’s Indust r ial melanism
(c) Past uer ’s Swann nech exp
(d) Ur ey miller s exp
70. Among t he following which is not pr er equisit e for nat ur al select ion-
(a) Unit capable of r epr oduct ion
(b) Occur r ence of her it able var iat ions or differ ences among unit s
(c) Pr esence of mor e t hen one such unit in sa me envir onment
(d) Envir onment al homeost asis
71. Which is false about st abilizing select ion-
(a) Const ant or unchanging envir onment
(b) Int r oduces het er ozygosit y
(c) Favour s aver age
(d) It t ends t o ar r est var iance & envir onment al changes
72. Indust r ial melanism is example of-
(a) Dir ect ional select ion (b) St abilizing Select ion
(c) Cyclic select ion (d) Disr upt ive Select ion
73. Appear ance of abnor mal char act er s which wer e nor mal in ot her supposedly ancest r al
r aces for example appear ance of cer vical fist ula in man which act ually cor r esponds t o
gill slit s ar e-
(a) Race at avism (b) Family at avism
(c) At avism of t er at ology (d) Ont ogenic at avism
74. The ascidian lar va on met amor phosis changes int o degener at e adult . This is example
of-
(a) Pr ogr essive met amor phosis (b) Dir ect ional
(c) Ret r ogr essive (d) Cyclic
75. C
14
is used t o det er mine age of fossils up t o-
(a) 10000 year s old (b) 25000
(c) 45000 (d) 1 lakh year
76. The clock of r ock is usely det er mined by-
(a) U
238
(b) K
40
(c) Bot h a & c (d) C
14
77. The t r ansit ional fossil for m which shows char act er ist ics of t wo differ ent gr oups of
living or ganisms is called as-
(a) Missing link (b) Connect ing link
(c) Living fossil (d) Link species
Practice Test Paper–Evolutionary Biology 295
78. Monot r emes (egg laying mammals) and mar supials (pouched mammals) ar e r est r ict ed
t o dist r ibut ion in-
(a) Asia (b) Aust r alia
(c) Afr ica (d) N. Amer ica
79. Among t he following hor mones, which ar e pr esent in all animals-
(a) Pepsin, t r ypsin (b) Tr ypsin, Amylase
(c) Pepsin, amylase (d) Tr ypsin, secr et in
80. Among t he following which is most conser ved in all eukar yot es and act as r espir at or y
pigment and accept s elect r on for m H
+
-
(a) Fer r odoxin (b) Cyt chr ome-c
(c) Cyct ochr ome c oxidase (d) NAD dehydr ogenase
81. Which hor mone is ident ical and int er changeable and pr esent in all animals is-
(a) Thyr oid (b) Adr enalin
(c) Somat ot r opin (d) Insulin
82. Ar t ificial select ion t ends t o-
(a) Change gene fr equency (b) Decr ease biodiver sit y
(c) Incr ease vigor (d) All of t he above
83. Pr okar yot es called ____ ar e similar in many ways t o eukar yot ic or ganisms.
(a) ar chae (b) pr ot ozoa
(c) Cyanobact er ia (d) dinoflagellat es
84. Among t he following which hor mone is almost similar in beef, sheep, pig, whale,
hor se, r abit and differ s only in one t o t hr ee amino acid posit ion-
(a) Glucagon (b) Insulin
(c) Somat ot r opin (d) Somat ost anin
85. The or der of following or ganisms as phylogenet ic closeness t o human-
(a) Chimpanzee > languor > lemur > t r ee shr ews
(b) Languor > chimpanzee > t r ee shr ews > lemur
(c) Chimpanzee > languor > t r ee shr ews > Lemur
(d) Languor > lemur > chimpanzee > t r ee shr ews
86. The gr adual mode of speciat ion in single lineage in which species diver ge in spur t s of
r elat ively r apid change which r esult in incr ease in species is t er med as-
(a) Punct uat ed equilibr ium (b) Adapt ive r adiat ion
(c) Anagenesis (d) Cladogenesis
87. Among t he following which st at ement is false about r -select ed species-
(a) Long gener at ion t ime (b) Lar ge number of offspr ing
(c) Shor t life cycle (d) Tendency t o disper se
296 CSIR-NET Life Sciences
88. When an r adioact ive phosphor us is incor por at ed int o DNA st r and, t he phospho diest er
bond gener ally br eak aft er a shor t while because-
(a) The bond number of phosphor us decr eases
(b) Phosphor us on r adioact ive decay conver t s int o sulphur , which has valency of
t wo
(c) It at t acks glycosidic bond
(d) It br eaks hydr ogen bond
89. Among t he following which is gener ally not ut ilized for est ablishing phylogenet ic
r elat ionship bet ween t wo species-
(a) % similar it y (b) Geogr aphical dist ance
(c) Mar coli dist ance (d) Amino acid sequence
90. Mangr oves ar e highly pr oduct ive ecosyst em but t hey ar e poor in bir d diver sit y because-
(a) Lack of st r uct ur al diver sit y
(b) Lack of food diver sit y
(c) Mor e number of pr edat or s t hat feed on bir ds
(d) Lack of br eeding place
91. Among t he following which would lead int o new species for mat ion-
(a) Incr eased r esour ces
(b) Niche over lapping t oler ance
(c) Niche specializat ion
(d) Lack of compet it ion
92. Or igin of life is not possible under pr esent envir onment al condit ions because-
(a) Hydr ogen is absent
(b) Pr esence of oxygen
(c) Lack of sour ce of ener gy
(d) Lack of r aw mat er ial for or igin of life
93. Disast er such as ear t hquake or fir e may r educe t he size of populat ion dr ast ically and
t he genet ic make up of t he small sur viving populat ion is unlikely t o be r epr esent at ive
of make up of or iginal populat ion t he sit uat ion is t er med as-
(a) Bot t le neck effect (b) Adapt ive r adiat ion
(c) Founder effect (d) Gene migr at ion
94. A cer t ain or ganism developed an adapt at ion as t o adapt exist ing envir onment but if
t his adapt at ion pr ove useful for some ot her funct ion as well, it is t er med as-
(a) Co-adapt at ion (b) Adapt ive r adiat ion
(c) Exadapt at ion (d) Co-evolut ion
95. In a populat ion fr equency of a homozygous r ecessive disease is 16% t hen t he fr equency
of dominant allele would be-
(a) 0.84 (b) 0.6
(c) 0.16 (d) 0.4
Practice Test Paper–Evolutionary Biology 297
96. In a populat ion individuals having het er ozygous phenot ype ar e mor e favor ed t hen
homozygous dominant which ar e mor e favor ed t hen homozygous r ecessive genot ypes,
under such condit ion-
(a) Recessive alleles would be lost fr om populat ion
(b) Dominant alleles would be lost
(c) Bot h alleles would r emain in populat ion
(d) Alleles would be lost r andomly
97. A gr oup of species which ar e phylogenit ically closer but t hey ar e lacking common
ancest or . Such an gr oup is r egar ded as-
(a) Monophylet ic (b) Polyphylet ic
(c) Par aphylet ic (d) Sympat r ic
98. Aft er implicat ion of Gr een Air Act in England which species become vir t ually absent -
(a) Bist on bet ular ia car bonifer a (b) Bist on bet ular ia t ypica
(c) Dr osophila (d) Apes Amer icana
99. If an allele is linked t o t he second allele at ot her locus which is favor ably select ed ar e
inher it ed t oget her . Such an movement of an allele wit hout any evolut ionar y benefit
t o next gener at ion is t er med as-
(a) Select ive dr ive (b) Evolut ionar y dr ive
(c) Hit ch hiking (d) Linkage
100. The mor phological modificat ion is t r ansfer r ed t o next gener at ion wit hout any pr esent
applicat ion which may pr ove beneficial in changed envir onment is t er med as –
(a) Exadapt at ion (b) Pr e-adapt at ion
(c) Analogous (d) Par alogous
Practice Test Paper-II
1. In t he fir st or der chemical r eact ion, if t he t ime t aken for half t he r eact ion t o be
consumed is t , t he t ime t aken for t hr ee-four t h t o be consumed is-
(a) 1.5 t (b) ½ t
(c) 2 t (d) 1.33 t
2. Which of t he following or ganism is most suit able t o ser ve as a bio-fer t ilizer ?
(a) Chlor ella (b) Agr obact er ia
(c) Azolla (d) Fr ankia
3. A r adioact ive compound Cs
137
wa s collect ed on 1
st
Febr uar y, 2002, and kept in a
sealed t ube. On t he 1
st
of J uly, 2002, it was found t hat only 3.125 % r adioact ivit y was
left . This means t hat t he half-life per iod of t he isot ope is
(a) 37.5 days (b) 30 days
(c) 25 days (d) 50 days
4. The t wo component of lichen is-
(a) Fungus and bact er ia (b) Fungus and br yophyt e
(c) Fungus and algae (d) Algae and Bact er ia
5. Car bon-14 under goes b-decay upon which it is conver t ed int o a new element having-
(a) Incr eased at omic number (b) Decr eased at omic number
(c) Incr eased at omic mass (d) Decr eased at omic mass
6. The fir st or ganisms t hat or iginat ed on ear t h wer e-
(a) Chemolit hot r ophs (b) Bact er iophages
(c) Phot oaut ot r ophic bact er ia (d) Uninucleat e eukar yot es
7. Ent amoeba hist olyt ica is spr ead by-
(a) Bit e of sand fly
(b) Bit e of anopheles mosquit o
(c) Consuming cont aminat ed food & wat er
(d) Blood t r ansfusion
8. If t he pH of t he solut ion is 9, t hen
(a) The solut ion is said t o be acidic
(b) The hydr oxyl ion concent r at ion is 10
–9
M
(c) The solut ion has t hr ee t imes mor e H
+
ion t han t he solut ion of pH 12
(d) The Hydr ogen ion concent r at ion is 10
–9
M
9. Half life of r adioact ive I
131
is 8 days. If you have 100 micr omoles of t his isot ope at one
inst ant , how many micr omoles will r emain at t he end of sixt een days?
(a) Zer o (b) 25
(c) 33.33 (d) 50
Practice Test Paper–Evolutionary Biology 299
10. Which of t he following will not ionize t he gas?
(a) X-r ays (b) a-r ays
(c) Elect r ons (d) g-r ays
11. Which of t he following is t he fossil for m?
(a) Ar cheopt er yx (b) Amphioxus
(c) Or nit hor hynchus (d) Per ipat us
12. Mammals or iginat ed fr om-
(a) Fishes (b) Amphibians
(c) Rept iles (d) Birds
13. “Recapit ulat ion t heor y” in evolut ion was pr oposed by-
(a) Ha eckel (b) Dar win
(c) Lamar ck (d) Cuvier
14. The half life of t he fir st or der r eact ion is
(a) Const ant & dependent on t he init ial concent r at ion of t he r eact ant
(b) Const ant & independent on t he init ial concent r at ion of t he r eact ant
(c) Not const ant & independent on t he init ial concent r at ion of t he r eact ant
(d) Dependent on t he t emper at ur e and pr essur e
15. One of t he following is t he char act er ist ics feat ur es of t he Aves-
(a) Four chamber ed hear t (b) Absence of nucleus in RBC
(c) Pneuma t ic bones (d) Abilit y t o fly
16. In t he life cycle of malar ial par asit e (Plasmodium vivax ), t he pr e-er yt hr ocyt ic cycle
occur s in-
(a) Liver (b) Bone ma r r ow
(c) Blood plasma (d) Kidney
17. Wher e would you place sponges among following-
(a) Par azoa (b) Met azoa
(c) Pr ot ozoa (d) Heliozoa
18. Evolut ion:
(a) is a r ever sible pr ocess
(b) occur s t hr ough var iat ions ar ising fr om changes in genet ic mat er ial
(c) is a fast pr ocess
(d) cannot t ake place any mor e because of oxidizing envir onment
19. Cr op r ot at ion for r est or ing soil fer t ilit y r efer s t o gr owing one of t he following cr ops
bet ween cer eal cr ops-
(a) Pulse cr ops (b) Gr ass
(c) Composit e (d) Tr ees
300 CSIR-NET Life Sciences
20. Kala-azar is caused by:
(a) Leishmania donovani (b) Trypanosoma gambiense
(c) Ent amoeba hist olyt ica (d) Plasmodium falciparum
21. The oldest micr ofossils discover ed so far of age 3.3 t o 3.5 billions year ago was-
(a) Coa cer va t es (b) Eobiont s
(c) Micr oshper es (d) Cyanobact er ia
22. In his book, “ The or igin of life (1938)” opar in submit t ed abiogenesis fir st but biogenesis
ever since”, t his t heor y is names as-
(a) Spont aneous gener at ion (b) Chemical or igin
(c) Pr imar y abiogenesis (d) Biogenenesis
23. Exper iment al evidence for molecular evolut ion of life was pr ovided by-
(a) Opar in (b) Haldane
(c) Ur ey & Miller (d) Sydney fox
24. Dur ing pr e-biot ic ear t h of life which chemical played impor t ant r ole in for mat ion of
nucleot ides-
(a) Met hane (b) Ammonia
(c) Hydr ogen (d) HCN
25. Which of t he following have lipid bilayer and can r epr oduce by budding-
(a) Coa cer va t es (b) Micr osper es
(c) Na nospher es (d) Pr ot obiont s
26. Which of t hem is evidence of evolut ion fr om biogeogr aphy as nar r at ed by Dar win-
(a) Embr yo devp. (b) Plat e t ect onics
(c) Dar win finches (d) Dar win t ur t les
27. Placent al mammals such as mouse wolf and Aust r alian mar supials such as mar supial
mouse, Tasmanian wolf shows-
(a) Par allel evolut ion (b) Conver gent
(c) Diver gent (d) Phyllet ic
28. Thor n of Bouganwalia and t endr ils of cucur bit s ar e-
(a) Homologous or ga n (b) Par alogous
(c) Analogous (d) Or t hologous
29. Which of t hem is not vest igial or gan in human-
(a) Ear muscles (b) Tail ver t ebr a
(c) Pr emolar (d) Appendix
30. The pr obable ener gy car r ier molecule dur ing init ial st age of or igin of life was-
(a) Amino acids (b) Nucleosides
(I) Nucleot ides (d) Sugar s
Practice Test Paper–Evolutionary Biology 301
31. Opar in and Sydney fox held t hat t he lar ge or ganic molecule synt hesized abiot ically
on pr imit ive ear t h for med lar ge colloidal aggr egat es due t o int er molecular int er act ions.
The colloidal par t icles wer e called-
(a) Micor ospher es (b) Eobiont s
(c) Ar chaebact er ia (d) Coa cer va t es
32. Pr obable t he fir st living for m was nut r it ionally-
(a) Chemoaut ot r ophs (b) Chemohet er ot r ophs
(c) Anoxygenic aut ot r ophs (d) Oxygenic aut ot r ophs
33. Which is most essent ial r equir ement for or igin of life on any planet -
(a) Wat er (b) Reducing envir onment
(c) Met hane (d) Nucleic acids
34. Seal flipper s, bir d and bat wings, hor se for e limbs and human ar ms ar e –
(a) Homologous or ga n (b) Analogaous or gan
(c) Vest igeal or gan (d) Or t hologous or gan
35. In bat t he wing is for med of fold of a int egument t er med as-
(a) Pt er odact yl (b) Pat agium
(c) Phalanges (d) Car pels
36. Evolut ion of new for ms in sever al dir ect ion fr om a common ancest or t ype is t er med
as –
(a) Phylogenic evolut ion (b) Anagenesis
(c) Cladogenesis (d) Adapt ive r adiat ion
37. The similar body shape bet ween animals of differ ent ly r elat ed gr oups r epr esent s-
(a) Conver gent evolut ion (b) Co-evolut ion
(c) Par allel evolut ion (d) All of t he above
38. Pr obable cause of appear ance of cer vical fist ula, t ail, hair on face in humans is-
(a) Mut at ion in homeot ic gene (b) r et r ogr essive evolut ion
(c) At avism (d) Genet ic dr ift
39. Adult fr og and ot her amphibian excr et es-
(a) Ammonia (b) Ur ea
(c) Ur ic acid (d) All
40. Ret r ogr essive met amor phosis is seen in-
(a) Amphibian t adpole (b) Ascidian lar vae
(c) Fishes (d) Rept iles
41. Galapagos island, t he biologist par adise, ar e sit uat ed on equat or about 960 Km west
of Ecuador in cont inent -
(a) Sout h Amer ica (b) Nor t h Amer ica
(c) Afr ica (d) Aust r alia
302 CSIR-NET Life Sciences
42. The Flor a and fauna of sout h Afr ica and west sout h Amer ica r esembles t o-
(a) Eur ope (b) Nor t h Amer ica
(c) Aust r alia (d) Asia
43. About 375 million year s ago, all pr esent cont inent s for med a single giant land mass
called pangea which split s in t wo islands masses Laur asia and Gondwana ar ound-
(a) 280 million year ago (b) 175 million year ago
(c) 120 million year ago (d) 80 million year ago
44. Among t he following which is an avian char act er -
(a) Homodent dent it ion
(b) Mat acar pel separ at e & car pomet acar pus absent
(c) V-shaped fer cula
(d) Eyes have scler ot ic ossicles
45. Radioact ive isot ope of car bon-14 can be used t o det er mine age of fossil up t o-
(a) 10,000 year old (b) 40,000 year old
(c) 1 Lakh year old (d) 10 lakh year old
46. When t he pr eser vat ion of genet ic var iabilit y is t hr ough het er ozygous super ior it y, it
is called-
(a) Het er opolymor phism (b) Balanced polymor phism
(c) St abilizing polymor phism (d) Dir ect ional polymor phism
47. Type of speciat ion due t o ploidy level is-
(a) Per ipat r ic (b) Par apat r ic
(c) Sympat r ic (d) Allopat ic
48. When t he t wo species ar e mor phologically almost ident ical but do not nor mally
int er br eed, such species ar e called-
(a) Taxonomic species (b) Sibling species
(c) Ecospecies (d) Ecospecies
49. Met hanogens ar e found in-
(a) Hot spr ings (b) Bior eact or s
(c) Flor a of cat t le r umen (d) All such places
50. Ar chaebact er ial cell wall is made up of-
(a) Lipo- polysacchar ide (b) Polysacchar ide and Pr ot eins
(c) Polysacchar ides only (d) Mur ien
51. Which of t hem can change t heir body shape-
(a) Vir uses (b) Bact er ia
(c) Mycoplasma (d) Eubact er ia
52. Afr ican sleeping sickness is caused by-
(a) Leishmania (b) Tr ypa nosoma
(c) Dict ost elluim (d) Physar ium
Practice Test Paper–Evolutionary Biology 303
53. The gamet ophyt e and spor ophyt e of br yophyt es ar e-
(a) Dependent on each ot her
(b) Independent t o each ot her
(c) Gamet ophyt e dependent on spor ophyt e
(d) Spor ophyt e dependent on gamet ophyt e
54. The chemical pr esent in flit is-
(a) Malat hion (b) DDT
(c) BHC (d) Aldicar b
55. Example of gr een manur e is-
(a) Rice (b) Sor ghum
(c) Maize (d) Sesbania
56. p and q in Har dy-Weinber g law r epr esent s-
(a) Allele fr equency (b) genot ype fr equency
(c) Het er ozygous fr equency (d) Tot al alleles in populat ion
57. Hybr id br eakdown occur s in-
(a) Donkey (b) Mule
(c) Liger (d) Hor se
58. The pr e-biot ic envir onment was-
(a) Oxidizing (b) Reducing
(c) No envir onment (d) Humid
59. Which chemical played impor t ant r ole in or igin of life-
(a) Met hane (b) Ammonia
(c) HCN (d) Hydr ogen
60. Ar t ificial select ion t ends t o-
(a) Change gene fr equency (b) Decr ease biodiver sit y
(c) Incr ease vigour (d) All t he above
61. Unifor m peeling away of soil sur face by t he act ion of flowing wat er is t er med as-
(a) Slip er osion (b) Rill er osion
(c) Gully er osion (d) Sheet er osion
62. Bacillus thuringenesis is used t o cont r ol-
(a) Bact er ial pat hogens (b) Fungal pat hogens
(c) Nemat ode (d) Insect pest s
63. In t he gr assland ecosyst em, t r ees donot r eplace t he gr asses as par t of ecological
succession because-
(a) Insect and fungi
(b) Limit ed sunlight & nut r ient s
(c) Wat er limit & r egular fir e or over gr azing
(d) Cool t emper at ur e
304 CSIR-NET Life Sciences
64. Nit r ogen fixat ion occur s in-
(a) Some bact er ia, cyanobact er ia & legumes
(b) Legumes and some bact er ia
(c) Some her baceous plant s & legumes
(d) All gr een plant s
65. The evolut ion of numer ous species in a shor t per iod of t ime fr om a single ancest r al
populat ion, such as Dar win’s finches, is called
(a) adapt ive r adiat ion (b) sympat r ic speciat ion
(c) gr adualism (d) nondisjunct ion
66. Mass ext inct ions t hat occur r ed in t he past
(a) cut t he number of species t o t he few sur vivor s left t oday
(b) r esult ed only fr om t he mer ging of t he cont inent s
(c) wer e followed by diver sificat ion of t he sur vivor s
(d) wiped out land animals, but had lit t le effect on mar ine life
67. What evidence most st r ongly suggest s t hat an impact by an ast er oid or met eor it e
may have caused t he ext inct ion of t he dinosaur s?
(a) Fossils show t hat dinosaur s suffer ed fr om cold and st ar vat ion
(b) Sediment ar y r ocks cont ain a layer of ir idium, a miner al t hat is uncommon on
Ear t h
(c) Ther e have been sever al near misses in r ecent year s
(d) The dinosaur s disappear ed r at her abr upt ly, vir t ually over night
68. Which of t he following would cast doubt on t he ast er oid-impact hypot hesis for t he
ext inct ion of t he dinosaur s?
(a) finding a cr at er 200 million year s old
(b) finding fossil dinosaur bones beneat h a layer of ir idium
(c) det er mining t hat bir ds ar e closely r elat ed t o dinosaur s
(d) finding fossil dinosaur bones above a layer of ir idium
69. Animals t hat possess homologous st r uct ur es pr obably
(a) ar e headed for ext inct ion
(b) evolved fr om t he same ancest or
(c) have incr eased genet ic diver sit y
(d) ar e not r elat ed
70. The wings of bir ds and insect s have t he same funct ion, but t hey do not have t he same
evolut ionar y or igin. Bir d and insect wings ar e
(a) homologous (b) phylogenet ic
(c) analogous (d) binomial
71. Which of t he following would be least useful in det er mining t he r elat ionships among
var ious species of or ganisms?
(a) DNA base sequences (b) homologous st r uct ur es
(c) fossils (d) amino acid sequences of pr ot eins
Practice Test Paper–Evolutionary Biology 305
72. A phylogenet ic t r ee of bir d families const r uct ed by cladist ic analysis would most clear ly
show which of t he following?
(a) char act er ist ics shar ed by all bir d families
(b) evolut ionar y r elat ionships among families
(c) families t hat look most alike
(d) analogous st r uct ur es shar ed by var ious species
73. Using cladist ic analysis, a t axonomist wishes t o const r uct a phylogenet ic t r ee showing
t he r elat ionships among var ious species of mammals. Dat a about which of t he following
would be least useful for t his pur pose?
(a) descr ipt ions of var ious t ypes of limbs (wings, legs, flipper s, et c.)
(b) dat a about skull bones
(c) t he fact t hat t eet h var y among differ ent t ypes of mammals
(d) DNA base sequences
74. Which of t he following was pr obably not pr esent in lar ge amount s in t he at mospher e
at t he t ime life is t hought t o have or iginat ed?
(a) wat er (H
2
O) (b) met hane (CH
4
)
(c) ammonia (NH
3
) (d) oxygen (O
2
)
75. Biologist s ar e int er est ed in t he r ole of clay in t he or igin of life. They t hink clay might
have
(a) supplied t he r aw mat er ials for or ganic compounds
(b) cat alyzed t he for mat ion of or ganic polymer s such as pr ot eins and RNA
(c) for med pr imit ive cell membr anes t hat could gr ow and divide
(d) cat alyzed t he for mat ion of monomer s such as amino acids and sugar s
76. Which of t he following is t hought t o have been t he fir st st ep in t he or igin of life?
(a) for mat ion of polypept ide spher es (b) for mat ion of or ganic monomer s
(c) r eplicat ion of pr imit ive genes (d) for mat ion of or ganic polymer s
77. Most bact er ia
(a) obt ain ener gy fr om sunlight and car bon fr om or ganic compounds
(b) obt ain bot h ener gy and car bon fr om inor ganic compounds
(c) obt ain ener gy fr om inor ganic compounds and car bon fr om CO
2
(d) obt ain bot h ener gy and car bon fr om or ganic compounds
78. Ener gy met abolism may have evolved when ear ly pr okar yot es
(a) used up t he oxygen in t heir envir onment
(b) began pr oducing poisonous wast e pr oduct s
(c) used up t he ATP in t heir envir onment
(d) began pr oducing significant amount s of oxygen
79. Unt il about 500 million year s ago, all living t hings wer e
(a) asexual (b) aut ot r ophic
(c) aquat ic (d) pr okar yot ic
306 CSIR-NET Life Sciences
80. You set your t ime machine for 3 billion year s ago and push t he st ar t but t on. When
t he dust clear s, you look out t he window. Which of t he following descr ibes what you
would pr obably see?
(a) plant s and animals ver y differ ent fr om t hose alive t oday
(b) a cloud of gas and dust in space
(c) gr een scum in t he wat er
(d) land and wat er st er ile and devoid of life
81. Select ion of Afr ican t r ibes is mor e of het er ozygous gene for RBC is due t o-
(a) Sever e malar ia
(b) Envir onment inst abilit y
(c) Mor e cases of Sickle cell anaemia
(d) bot h a & c
82. Molecular clock of evolut ion could be t r aced on basis of-
(a) Compar ison of Shor t ar m of 16-S RNA
(b) Subst it ut ion in amoniacids of polypept ide due t o mut at ion
(c) DNA finger pr int ing
(d) Fossil st udy
83. Among t he following t he pr ogenit or s of mammals wer e-
(a) Aves (b) Pisces
(c) Amphibians (d) Rept iles
84. Abr upt change in gene fr equency in isolat ed populat ion is t er med as-
(a) Adapt ive r adiat ion (b) Allopat r ic speciat ion
(c) Random dr ift (d) Mut at ion
85. Main cause of loss of Biodiver sit y is-
(a) Pollut ion (b) Populat ion explosion
(c) Habit at dest r uct ion (d) Over exploit at ion
86. Which r adioisot ope is used for est imat ing age of r ocks-
(a) U
238
& K
40
(b) U
235
& C
14
(c) U
238
& H
3
(d) C
14
& H
3
87. What is main cause of evolut ion of new species-
(a) Nat ur al select ion (b) Compet it ion
(c) Mut at ion (d) Hybr idizat ion
88. Phylogenet ic r elat ionship can be mor e pr ecisely est ablished by compar ing-
(a) Amino acid sequence (b) DNA
(c) r -RNA (d) m-RNA
89. In a sample fr om an Afr ican populat ion, t he fr equency of L
M
and L
N
alleles wer e
0.78 and 0.22 r esp. What ar e expect ed fr equency of MN phenot ypes-
(a) 0.8 (b) 0.02
(c) 0.34 (d) 0.016
Practice Test Paper–Evolutionary Biology 307
90. Phylogenet ic r elat ionship in plant s can be best est ablished by-
(a) Allozymes (b) Alkaloids
(c) I soenzymes (d) Mor phology
91. The Cuvier ’s t heor y, which st at es t hat t her e had been sever al cr eat ions, each pr eceded
by anot her due t o some geogr aphical dist ur bances. Such t heor y is t er med as-
(a) Theor y of cat ast r ophism (b) Special cr eat ion
(c) Pangenesis (d) Et er nit y of life
92. Whih is cor r ect mat ched among following connect ing links-
(a) Euglena- Rept iles and mammals
(b) Pr ot er ospongia- Car t ilaginous and bony fishes
(c) Per ipat us- Annelida & Mollusca
(d) Neoplina- Annelida & Ar t hopoda
(e) Balaonglossus- Chor dat e and Non-chor dat e
(f) Chimer a- Fishes and Amphibians
(g) Lung fishes- Pr ot ozoa & por ifer a
(h) Pr ot ot her ia- Plant s and animals
93. Ancient cyanobact er ia, found in fossil st r omat olit es, wer e ver y impor t ant in t he hist or y
of life because t hey
(a) wer e pr obably t he fir st living t hings t o exist on Ear t h
(b) pr oduced t he oxygen in t he at mospher e
(c) ar e t he oldest known ar chae
(d) wer e t he fir st mult icellular or ganisms
94. In t er ms of nut r it ion, aut ot r ophs ar e t o het er ot r ophs as
(a) algae ar e t o slime molds
(b) ar chae ar e t o bact er ia
(c) slime molds ar e t o algae
(d) kelp ar e t o diat oms
95. The bact er ia t hat cause t et anus can be killed only by pr olonged heat ing at t emper at ur es
consider ably above boiling. This suggest s t hat t et anus bact er ia
(a) have cell walls cont aining pept idoglycan
(b) pr ot ect t hemselves by secr et ing ant ibiot ics
(c) secr et e endot oxins
(d) pr oduce endospor es
96. Bact er ia such as S t rept ococcus pneumoniae can cause disease in humans when
(a) t he host s immune syst em is compr omised
(b) t he bact er ium develops r esist ance t o all ant ibiot ics
(c) a co-host such as St aphylococcus is pr esent on t he host
(d) t hat pr ot ozoan has pat hogenic fact or s such as pili or capsules
308 CSIR-NET Life Sciences
97. Gr am-negat ive bact er ia have ________ pept idoglycan t han Gr am-posit ive cells and
t heir cell walls ar e _________ complex st r uct ur ally.
(a) mor e . . . mor e
(b) mor e . . . less
(c) less . . . less
(d) less . . . mor e
98. Genes for t he r esist ance of ant ibiot ics ar e usually locat ed
(a) on t he ma in chr omosome
(b) in mit ochondr ia
(c) in t he cell wall
(d) on plasmids
99. Bact er ia t hat can conver t at mospher ic nit r ogen int o ammonia ar e called
(a) nit r ogen r educer s
(b) nit r ogen fixer s
(c) nit r ogen oxidizer s
(d) nit r ogen het er ot r ophs
100. Humans ar e phlyogenet ically mor e close t o-
(a) Monkeys
(b) Tr ee shr ews
(c) Lemur s
(d) Sinipsids
6
Environmental Biology
MAJOR COMPONENTS OF ECOLOGY
1. Ecology
(a) “The scient ific st udy of t he int er act ions bet ween or ganisms and t heir envir onment s
is called ecology.”
(b) “Ecology concer ns it self wit h t he int er r elat ionships of living or ganisms, plant or animal,
and t heir envir onment s; t hese ar e st udied wit h a view t o discover ing t he pr inciples
which gover n t he r elat ionships.
2. Environment
(a) An or ganism’s envir onment may be dist inguished int o an abiot ic component and a
biot ic component .
(b) “Or ganisms ar e affect ed by t heir envir onment but , by t heir ver y pr esence and act ivit ies,
t hey also change it —oft en dr amat ically.”
3. Abiotic component
(a) The abiot ic component of an envir onment ar e all of t he non-living component s of an
or ganism’s envir onment .
(b) These include such t hings as Temper at ur e; Light ; Wat er ; Wind; Nut r ient s; Subst r at e
(e.g., r ock and soil); Per iodic dist ur bances.
(c) Gener ally, a given or ganism is capable of sur viving over only a limit ed r ange of
abiot ic var iables, and t he envir onment s in which one (or mor e) abiot ic component
r anges out side of an or ganism’s r ange of t oler ance will not be able t o suppor t st able
populat ions of t hat or ganism.
310 CSIR-NET Life Sciences
4. Biotic component
(a) The biot ic component of an envir onment ar e all of t he ot her or ganisms found in an
envir onment wit h which an or ganism makes cont act , dir ect ly or indir ect ly. These
or ganisms may be compet ing, pr eying upon, being pr eyed upon, pr oviding shelt er , or
in some ot her way impact on t he envir onment .
5. Evolution in real time
(a) The complexit y associat ed wit h ecology explains in par t why t he st udy of evolut ion is
so int ensely difficult : Evolut ion happens wit hin a cont ext of ecology, i.e., in r eal
ecosyst ems, one or ganism at a t ime.
(b) Par t of t he complexit y associat ed wit h ecology, however , is a consequence of t he
impact of evolut ion on ecosyst ems: Ecosyst ems ar e not only t he pr oduct s of evolut ion,
t hey also cont ain populat ions t hat ar e act ively evolving, all of t he t ime.
(c) In ot her wor ds, ecology is essent ially evolut ion r unning in r eal t ime, while evolut ion
is essent ially t he pr oduct of enor mous number s of ecological int er act ions bet ween
or ganisms and t heir biot ic and abiot ic envir onment s.
6. Principle of allocation
(a) One way t o under st and ecology is in t er ms of flows of ener gy; or ganisms t ake in
ener gy and t hen use t hat ener gy t o sur vive and t o r epr oduce. All adapt at ions ar e
compr omises, no or ganism is per fect ly adapt ed t o ever yt hing, and ever yt hing cost s
ener gy. An or ganism must balance out it s allocat ion of ener gy t o sur vival and it s
need t o allocat e ener gy t o r epr oduct ion.
(b) Genot ypes t hat st r ike a good balance bet ween allocat ion t o sur vival and t o r epr oduct ion,
such t hat net r epr oduct ion is lar ge compar ed wit h ot her genot ypes, ar e said t o have
higher r elat ive fit nesses. These ideas for m t he basis of t he pr inciple of allocat ion.
(c) “Each or ganism has a limit ed amount of ener gy t hat can be allocat ed for obt aining
nut r ient s, escaping fr om pr edat or s, coping wit h envir onment al fluct uat ions, gr owt h
and r epr oduct ion.”
(d) Ener gy allocat ed t o sur vival is not available for r epr oduct ion.
(e) “Complex life like animals and plant s needs a lot of ener gy.” Relat ively simple
or ganisms have lower ener gy needs, but t end also t o be mor e limit ed in wher e t hey
live or how much ener gy t hey can obt ain per unit t ime.
7. Adaptation
(a) Or ganisms can r espond t o var iat ions in t he envir onment wit h var iet y of adapt at ions.
(i) Behaviour al adapt at ions ar e almost inst ant aneous in t heir effect s and easily
r ever sed, wher eas Physiological adapt at ions may be implement ed and changed
over t ime scales r anging fr om seconds t o weeks.
(ii) Mor phological adapt at ions may develop over t he lifet imes of individual or ganisms
or bet ween gener at ions.
(iii) Adapt ive genet ic changes in populat ions ar e slower st ill, usually evolving over
sever al gener at ions.
Environmental Biology 311
(b) The appr opr iat e r esponse t o envir onment al change depends on t he dur at ion of t hat
change.”
(c) “The dist inct ion bet ween shor t -t er m adjust ment s on t he scale of ecological t ime and
adapt at ion on t he scale of evolut ionar y t ime begins t o blur when we consider t hat t he
r ange of r esponses of an individual t o changes in t he envir onment is it self t he pr oduct
of evolut iona r y hist or y.” Tha t is, “p h e n ot yp i c p l a s t i ci t y” is it self a pr oduct of
evolut ion
(d) “In gener al, plant s ar e mor e mor phologically plast ic t han animals; t his r esponse helps
t hem compensat e for t heir inabilit y t o move fr om one envir onment al pat ch t o anot her .”
MAJOR AREAS OF ECOLOGICAL STUDY
8. Hierarchical study of ecology
(a) The st udy of ecology is oft en achieved by concent r at ing on a cer t ain level of a hier ar chy
of ecological st udy, just as mor e r educt ionist for ms of biology focus on cer t ain levels
of or ganisms r anging fr om t he molecular , t hr ough t he cellular , t hr ough t he or ganismal
(b) “Ecology ult ima t ely dea ls wit h t he highest levels in t he hier a r chy of biologica l
or ganizat ion. The web of int er act ions at t he hear t of ecological phenomena is what
makes t his br anch of biology so engaging.”
(c) The hier ar chies in ecological st udy include
(i) Or ganismal ecology
(ii) Populat ion ecology
(iii) Communit y ecology
(iv) Ecosyst em ecology
9. Organismal ecology
(a) Or ganismal ecology is an at t empt t o under st and how t he char act er ist ics of individual
or ganisms impact on t he abilit y of t hose or ganisms t o int er act wit h t heir envir onment .
(b) Thus, for example, infer r ing t hat gir affes use t heir long necks t o r each leaves found
high in t r ees is an example of or ganismal ecology (t hough oft en infer ences ar e just a
bit mor e subt le and difficult t han t his example).
10. Behavioral ecology
(a) Essent ially a subset of or ganismal ecology, behaviour al ecology st udies t he non-
physiological, non-mophological/anat omical adapt at ions or ganisms possess, and t he
impact t hose adapt at ions have on t he sur vival and r epr oduct ion of or ganisms.
(b) Behaviour , in ot her wor ds, is how or ganisms act , and differ ent behavior s can have
differ ent impact s on t he Dar winian fit ness of or ganisms.
11. Population ecology
(a) Populat ion ecology is t he st udy of t he size and composit ion of populat ions of or ganisms.
(b) An example of populat ion ecology would be t he st udy of t he fact or s which influence
t he car r ying capacit y of a given envir onment , i.e., t he number of individuals an
envir onment can st ably sust ain.
312 CSIR-NET Life Sciences
12. Community ecology
(a) A communit y is t he assemblage of differ ent species of or ganisms wit hin a given
envir onment . Communit y ecology is t he st udy of t he int er act ions bet ween t hese
or ganisms, e.g., pr edat ion, par asit ism, compet it ion, et c.
13. Ecosystem ecology
(a) An ecosyst em is t he assemblage of t he biot ic and abiot ic component s of a given
envir onment . Oft en ecosyst ems ar e r easonably unambiguously defined (a lake, a
for est , et c). Under st a nding even a n a ppr oxima t ion of wha t goes on wit hin a n
ecosyst em, any ecosyst em, can be an over whelming challenge.
(b) The ever glades ecosyst em as a funct ion of alt it ude and ot her fact or s.
14. Landscape ecology
(a) “Looking beyond t he four basic levels of ecology, we come t o landscape ecology, which
deals wit h ar r ays of ecosyt ems and how t hey ar e ar r anged in a geogr aphic r egion. A
landscape or seascape consist s of sever al differ ent ecosyst ems linked by exhanges of
ener gy, mat er ials, and or ganisms. The landscape level of r esear ch focuses on t he
ways in which int er act ions among populat ions, communit ies, and ecosyst ems ar e
affect ed by t he juxt aposit ion of differ ent ecosyst ems, such as st r eams, lakes, old-
gr owt h for est s, and t he for est pat ches t hat have had t heir t r ees r emoved by clear -cut
logging.”
15. Maintaining homeostasis
(a) Pa r t of t he ener gy expended on sur viva l goes t owa r d ma int a ining t he int er na l
50
100 150 200 250
100
80
60
40
20
Exponential vs Logistic
Environmental
Resistance
Carrying Capacity (K)
Pop.
size
Time
Environmental Biology 313
envir onment of an or ganism. The act ive maint enance of t he int er nal envir onment of
an or ganism (by t he or ganism) is t er med h ome os t a s i s .
(b) Some or ganisms spend consider ably mor e ener gy on maint aining t heir int er nal
envir onment wit hin r elat ively nar r ow const r aint s (r e gu l a t or s ).
(c) Ot her spend less ener gy on maint aining t heir int er nal envir onment because t hey do
not const r ain it nar r owly (con for me r s ).
(d) Any ener gy not spent on homeost asis is pot ent ially available for ot her needs such as
r epr oduct ion (p r i n ci p l e of a l l oca t i on ).
(e) Not e t hat t his pr inciple for ms t he basis of t he conflict ing st r at egies of specializat ion
(e.g., confor mer s) ver sus gener alizat ion (e.g., r egulat or s).
(f) Specialist s pot ent ially have mor e ener gy available t o r epr oduce because t hey ver y
efficient ly obt ain ener gy necessar y for maint aining homeost asis, t hough t his advant age
is maint ained only so long as t he envir onment r emains amenable t o t he specialist ’s
needs (not e: t r y t o avoid equat ing t he concept of specializat ion wit h t he concept of
specializat ion wit h r espect t o for aging; t he lat t er , specializat ion wit h r egar d t o diet ,
is essent ially a subset of t he for mer ).
16. Conformers
(a) For or ganisms wit hin r elat ively st able envir onment s, ener gy can be made available
for ot her uses if int er nal envir onment s ar e allowed t o var y as ext er nal envir onment s
var y. Such or ganisms may be t er med confor mer s.
(b) Not e t hat a confor mer may be ver y successful wit hin it s r elat ively st able envir onment ,
but less able t o sur vive out side of t his envir onment .
(c) “Con for mer s t h a t li ve i n ver y st a ble en vi r on men t s . . . mi gh t be a ble t o ch a n n el
mor e en er gy i n t o gr owt h a n d r ep r od u ct i on . However , t h e i n t oler a n ce of su ch
s p e ci a l i s t s t o e n vi r on me n t a l ch a n ge s e ve r e l y r e s t r i ct s t h e i r ge ogr a p h i ca l
d i s t r i b u t i on .”
17. Regulators
(a) At t he opposit e end of t he spect r um ar e t he r egulat or s (i.e., ver sus confor mer s).
These or ganisms expend a gr eat deal of ener gy t o keep t heir int er nal envir onment
const ant r egar dless of t he nat ur e of t heir abiot ic envir onment .
(b) Such or ganisms may be mor e adapt able, but at t he cost of gr eat expendit ur es of
ener gy t hat could ot her wise be put t owar d such t hings as r epr oduct ion.
(c) “Re gu l a t or s t h a t a l l oca t e a l a r ge r fr a ct i on of t h e i r e n e r gy t o cop i n g wi t h
e n vi r on me n t a l ch a n ge s ma y gr ow a n d p r op a ga t e l e s s e ffi ci e n t l y, b u t s u ch
or ga n i s ms a r e a b l e t o s u r vi ve a n d r e p r od u ce ove r a wi d e r r a n ge of va r i a b l e
e n vi r on me n t s .”
18. Graininess
(a) Envir onment al gr ain r efer s t o t he pat chiness of an envir onment , and t he pat chiness
of an envir onment is per ceived differ ent ly by differ ent or ganisms. A coar sely gr ained
envir onment has pat ches which ar e lar ge enough t hat t hey may be dist inguished. A
finely gr ained envir onment has pat ches which ar e so small t hat t hey may not be
314 CSIR-NET Life Sciences
r eadily dist inguished, and an “or ganism may not even behave as t hough pat ches
exist ,” but what is fine-gr ained t o one (t ypically lar ger ) or ganism may by coar se-
gr ained t o anot her (t ypically smaller ) or ganism.
19. Population ecology
(a) Populat ion ecology st udies or ganisms fr om t he point of view of t he size and st r uct ur e
of t heir populat ions.
(b) A populat ion ecologist st udies t he int er act ion of or ganisms wit h t heir envir onment s
by mea sur ing pr oper t ies of popula t ions r a t her t ha n t he beha vior of individua l
or ganisms.
(c) Pr oper t ies of populat ions include
• Populat ion size (size)
• Populat ion densit y (densit y)
• Pat t er ns of disper sion (disper sion)
• Demogr aphics (demogr aphics)
• Populat ion gr owt h (gr owt h)
• Limit s on populat ion gr owt h (limit s)
(d) Not e t hat all of t hese pr oper t ies ar e not t hose of individual or ganisms but inst ead ar e
pr oper t ies which exist only if one consider s mor e t han one or ganism at any given
t ime, or over a per iod of t ime (i.e., t hey ar e emer gent pr oper t ies)
(e) “The char act er ist ics of a populat ion ar e shaped by t he int er act ions bet ween individuals
and t heir envir onment s on bot h ecological and evolut ionar y t ime scales, and nat ur al
select ion can modify t hese char act er ist ics in a populat ion.”
(f) Thus, populat ion ecology also goes beyond consider at ion of just populat ion par amet er s
and addit ionally consider s how t he char act er ist ics of individual or ganisms impact on
populat ion par amet er s.
POPULATION PROPERTIES
1. Population
(a) A populat ion in an ecological sense is a gr oup of or ganisms, of t he same species,
which r oughly occupy t he same geogr aphical ar ea at t he same t ime.
(b) Individual member s of t he same populat ion can eit her int er act dir ect ly, or may int er act
wit h t he disper sing pr ogeny of ot her member s of t he same populat ion (e.g., pollen).
Populat ion member s int er act wit h a similar envir onment and exper ience similar
envir onment al limit at ions.
2. Population size
(a) A populat ion’s size depends on how t he populat ion is defined. If a populat ion is defined
in t er ms of some degr ee of r epr oduct ive isolat ion, t hen t hat populat ion’s size is t he
size of it s gene pool.
(b) If a populat ion is defined in t er ms of some geogr aphical r ange, t hen t hat populat ion’s
size is t he number of individuals living in t he defined ar ea.
Environmental Biology 315
(c) Ecologist s t ypically ar e mor e concer ned wit h t he lat t er means of defining a populat ion
since t his is bot h easier t o do and is a mor e pr act ical measur e if one is int er est ed in
det er mining t he impact of a given populat ion on a given ecosyst em, or vice ver sa.
3. Population density
(a) Given t hat a populat ion is defined in t er ms of some nat ur al or ar bit r ar ily defined
geogr aphical r ange, t hen populat ion densit y may be defined as simply t he number of
individual or ganisms per unit ar ea.
(b) Differ ent species, of cour se, exist at differ ent densit ies in t heir envir onment s, and
t he same species may be able t o achieve one densit y in one envir onment and anot her
in a differ ent envir onment . Populat ion densit ies may addit ionally be det er mined in
t er ms of some measur e ot her t han populat ion size per unit ar ea such as populat ion
mass per unit ar ea.
4. Patterns of dispersion
(a) Individual member s of populat ions may be dist r ibut ed over a geogr aphical ar ea in a
number of differ ent ways including
• Clumped dist r ibut ion (at t r act ion)
• Unifor m dist r ibut ion (r epulsion)
• Random dist r ibut ion (minimal int er act ion/influence)
(b) Clumping may r esult eit her fr om individual or ganisms being at t r act ed t o each ot her ,
or individual or ganisms being at t r act ed mor e t o some pat ches wit hin a r ange t han
t hey ar e t o ot her pat ches; t he net effect is t hat some par t s of t he r ange will have a
lar ge number of individuals wher eas ot her s will cont ain few or none.
(c) A unifor m dist r ibut ion means t hat appr oximat ely t he same dist ance may be found
bet ween individual or ganisms; unifor m dist r ibut ions r esult fr om individual or ganisms
act ively r epelling each ot her .
(d) A r andom dist r ibut ion means t hat wher e individual or ganisms ar e found is only
minimally influenced by int er act ions wit h ot her member s of t he same populat ion,
and r andom dist r ibut ions ar e uncommon; “Random spacing occur s in t he absence of
st r ong at t r act ions or r epulsions among individuals of a populat ion.”
(e) Not e t hat bot h clumping and unifor m dist r ibut ions suggest t hat individual or ganisms
ar e eit her int er act ing wit h one anot her (act ively seeking each ot her out or act ively
avoiding each ot her ), or ar e all compet ing wit h one anot her for t he same limit ed
r esour ces, r egar dless of t he over all populat ion densit y (as in t he case of clumping
which r esult s fr om geogr aphical pat chiness).
DEMOGRAPHICS
5. Demographics
(a) A populat ion’s demogr aphics ar e it s vit al st at ist ics, par t icular ly t hose st at ist ics which
can impact on pr esent and fut ur e populat ion size.
(b) Two st at ist ics t hat ar e of par t icular impor t ar e a populat ion’s age st r uct ur e and a
populat ion’s sex r at io.
316 CSIR-NET Life Sciences
(c) Addit ional consider at ions (in human populat ions and for example) ar e consider ed t o
t he r ight ®
6. Age structure
(a) Age st r uct ur e r efer s t o t he size of cohor t s wit hin a populat ion.
(b) Par amet er s r elat ed t o age st r uct ur e include
(i) Fecundit y (bir t h r at e)
(ii) Gener at ion t ime
(iii) Deat h r at e
7. Cohort
(a) A cohor t is a gr oup of individuals all of whom have t he same age.
(b) In a t ypical populat ion, t he size of cohor t s will var y wit h age. For example, in a
t ypical populat ion, younger cohor t s will be lar ger (i.e., mor e individuals per cohor t )
t han older cohor t s, all else being equal.
8. Fecundity [birth rate]
(a) Fecundit y r efer s t o t he aver age bir t h r at e associat ed wit h a populat ion.
(b) The gr eat er a populat ion’s fecundit y, all else held const ant , t he fast er a populat ion
will incr ease in size. Not e t hat fecundit y t ypically var ies wit h t he age of individuals.
9. Generation time
(a) Gener at ion t ime is simply t he aver age span bet ween t he bir t h of individuals and t he
bir t h of t heir offspr ing.
(b) “Ot her fact or s being equal, a shor t er gener at ion t ime will r esult in fast er populat ion
gr owt h.”
(c) Not e t hat species which ar e capable of r epr oducing mor e t han once will display an
over lapping of gener at ions which basically means t hat par ent al cohor t s and pr ogeny
cohor t s can be alive (and pot ent ially compet ing wit h one anot her ) at t he same t ime.
Education
Work
background
Parental
status
Geographic
Location
Religious
belefs
Marital
status
Income
Race
Ethnicity
Sexual/
Affectual
Orientation
Physical
Abilities/
Qualities
Age
Gender
Environmental Biology 317
(d) Not e t hat anot her way of saying t his is t hat when life expect ancies exceed t he
minimum t ime bet ween gener at ions, gener at ions will over lap.
10. Death rate
(a) Deat h r at e is t he r at e at which individuals of a cer t ain age die.
(b) Not e t hat deat h r at es oft en var y wit h age wit h eit her t he ver y young or t he ver y old
displaying t he gr eat est deat h r at es.
(c) Not e addit ionally t hat populat ion gr owt h occur s when over all bir t h r at es exceed
over all deat h r at es.
11. Sex ratio
(a) Mor e oft en t han not t he r at e at which a populat ion may gr ow is dependent on t he sex
r at io in t he populat ion; t he fewer females, t he slower t he r at e of populat ion gr owt h.
(b) This, of cour se, is because ut er uses ar e limit ing and males oft en can inseminat e
mor e t han one female.
(d) This gener alizat ion falls apar t , however , when males ar e limit ed in t heir abilit y t o
inseminat e mor e t han one female, or when males cont r ibut e significant ly t o t he
r aising of offspr ing.
(e) Figur e shows sex r at ios (New Sout h Wales) as t hey var y wit h age (unit s on y axis ar e
in living males per 100 females):
SURVIVORSHIP CURVES
12. Survivorship curves
(a) Obser ving a ge st r uct ur e gr a phica lly ca n pr ovide insight s int o a species’ (or a
populat ion’s) ecology. Sur vivor ship cur ves gr aph cohor t size against r elat ive age.
(b) Se e F i gu r e , I d e a l i ze d s u r vi vor s h i p cu r ve s .
(c) The t ypical sur vivor ship cur ve shows cohor t size declining wit h age.
(d) Ther e exist t hr ee gener al t ypes of sur vivor ship cur ves
• Typ e I , I I a n d I I I
(e) Not e in t he following sur vivor ship cur ves t hat t he y axis is logar it hmic!!!
120
100
80
60
40
0-4 10-14 20-24 30-34 40-44 50-54 60-64 70-74 80-84
Actual sex ratio
Even sex ratio
Age Groups
318 CSIR-NET Life Sciences
Type I survivorship curves
(a) Because individuals t end t o die exponent ially due t o accident s or pr edat ion, it oft en is
a good st r at egy t o r epr oduce r elat ively ear ly in a life span r at her t han r elat ively lat e
(i) That way individuals achieve r epr oduct ion while t hey st ill have a r easonable
likelihood of being alive.
(ii) This is assuming, of cour se, t hat t he goal is a Dar winian one, i.e., maximizing
one’s r epr oduct ive out put .
(iii) Not e t hat how such a st r at egy wor ks is complicat ed if individual fecundit y incr eases
wit h age.
(b) Ver y oft en for a given species t her e will be some age at which individuals ar e maximally
fecund. Species t hat combine maximum fecundit y wit h ear ly ages t ypically do so at
t he expense of t heir abilit y t o sur vive long per iods (i.e., t his is an example of t he
pr inciple of allocat ion).
(c) A sur vivor ship cur ve of such individuals may display a r elat ively shallow slope while
individuals ar e younger (i.e., maximally r obust and maximally r epr oduct ive) but t hen
show an abr upt incr ease in deat h r at e at ages t hat ar e coincident t o declines in
fecundit y.
(d) Humans, of cour se, have a t ype I sur vivor ship cur ve; evolut ion makes us get mar r ied
young and have lot s of babies befor e a saber t oot hed t iger comes along and picks us
off, i.e., ®
Type II survivorship curves
(a) The simplest t ype of decline is exponent ial, i.e., t he deat h r at e for ever y cohor t is t he
same.
(b) These sur vivor ship cur ves gr aph as a st r aight line on semi-logar it hmic gr aph paper
(i.e., as pr esent ed in a t ypical sur vivor ship cur ve).
(c) The individuals in populat ions t hat display a t ype II cur ve ar e t hose t hat bot h do not
age and ar e bor n as fully fit as adult s, e.g., hydr a Individuals ar e lost in t hese
populat ions most ly t o accident s and pr edat ion.
Type III
(Oysters)
Type II
(Hydra, Songbirds)
1.0
0.1
0.01
0.001
Birth Maximum
Life Span
Age of Organism
(scaled to maximum life spon
for each species)
Type I (Man)
F
r
a
c
t
i
o
n

o
f

O
r
g
a
n
i
s
m
s

S
u
r
v
i
v
in
g
Environmental Biology 319
Type III survivorship curves
(a) The ot her side of t he sur vivor ship coin is t he degr ee of invest ment in individual
pr ogeny. Some or ganisms invest a gr eat deal in each offspr ing and t hose or ganisms
ar e (ideally at least ) r ewar ded wit h r elat ively high sur vivor ship at ear ly ages.
(b) Ot her or ganisms invest lit t le in individual offspr ing, and display ver y low ear ly-age
sur vivor ship (which t hey make up for by pr oducing bucket s of offspr ing).
(c) Or ganisms t hat pr oduce lar ge number s of cheap pr ogeny and which display minimal
declines in fecundit y wit h age, if t hey sur vive t heir yout h, display t ype III sur vivor ship
cur ves. Examples include sea t ur t les and t r ees.
(d) That is, t ype III sur vivor ship species have a ver y lar ge r at e of mor t alit y when young,
but should t hey sur vive t heir yout h, t hey put significant ener gy int o cont inued sur vival
since t he longer t hey sur vive, t he mor e pr ogeny t hey will pr oduce.
LIFE HISTORIES
13. Life history
(a) “The t r ait s t hat affect an or ganism’s schedule of r epr oduct ion and sur vival (fr om
bir t h t hr ough r epr oduct ion t o deat h) make up it s life hist or y.” The st udy of life hist or y
char act er ist ics is t he det ailed st udy of t hose ecological and evolut ionar y par amet er s
t hat impact on sur vivor ship cur ves.
(b) “In many cases t her e ar e t r ade-offs bet ween sur vival and t r ait s such as clut ch size
(number of offspr ing per r epr oduct ive episode), fr equency of r epr oduct ion, a nd
invest ment in pa r ent a l ca r e. The t r a it s t ha t a ffect a n or ga nism’s schedule of
r epr oduct ion and deat h make up it s life hist or y. Of cour se, a par t icular life hist or y
pat t er n, like most char act er ist ics of an or ganism, is t he r esult of nat ur al select ion
oper at ing over evolut ionar y t ime.”
(c) In ot her wor ds, t h e Da r wi n i a n goa l i s t o ma xi mi ze l i fe t i me r e p r od u ct i ve
ou t p u t , and t his can be achieved by having babies mor e r apidly or living longer , or
some combinat ion of t he t wo, as well as by var ying many addit ional det ails having t o
do wit h sur vival and r epr oduct ion.
320 CSIR-NET Life Sciences
14. Allocation of limited resources
(a) “Dar winian fit ness is measur ed not by how many offspr ing ar e pr oduced but by how
many sur vive t o pr oduce t heir own offspr ing: Her it able char act er ist ics of life hist or y
t hat r esult in t he most r epr oduct ively successful descendant s will become mor e common
wit hin t he populat ion. If we wer e t o const r uct a hypot het ical life hist or y t hat would
yield t he gr eat est lifet ime r epr oduct ive out put , we might imagine a populat ion of
individuals t hat begin r epr oducing at an ear ly age, have lar ge clut ch sizes, and
r epr oduce many t imes in a lifet ime. However , nat ur al select ion cannot maximize all
t hese var iables simult aneously, because or ganisms have a finit e ener gy budget t hat
mandat es t r ade-offs. For example, t he pr oduct ion of many offspr ing wit h lit t le chance
of sur vival may r esult in fewer offspr ing t hat can compet e vigor ously for limit ed
r esour ces in an alr eady dense populat ion.”
(b) “The life hist or y we obser ve in or ganisms r epr esent a r esolut ion of sever al conflict ing
demands. An impor t ant par t of t he st udy of life hist or ies has been under st anding t he
r elat ionship bet ween limit ed r esour ces and compet ing funct ions: Time, ener gy, and
nut r ient s t hat ar e used for one t hing cannot be used for somet hing else.”
(c) “These issues can be phr ased in t er ms of t hr ee basic quest ions:
(i) How oft en should an or ganism br eed?
(ii) When should it begin t o r epr oduce?
(iii) How many offspr ing should it pr oduce dur ing each r epr oduct ive episode?
(d) The way each populat ion r esolves t hese quest ions r esult s in t he int egr at ed life hist or y
pat t er ns we see in nat ur e.” (all one quot e st ar t ing wit h (c) but br oken up for clar it y)
(e) “Many life hist or y issues involve balancing t he pr ofit of immediat e invest ment in
offspr ing against t he cost t o fut ur e pr ospect s of sur vival and r epr oduct ion. These
issues can be summar ized by t hr ee basic “decisions”: when t o begin r epr oducing, how
oft en t o br eed, and how many offspr ing t o pr oduce dur ing each r epr oduct ive episode.
The var ious “choices” ar e int egr at ed int o t he life hist or y pat t er ns we see in nat ur e.”
15. Semelparity (big bang)
(a) Or ganisms t hat pr oduce one clut ch of offspr ing (pr ogeny) per life t ime ar e said t o be
semelpar ous (i.e., t o display semelpar it y). The advant age of semelpar it y is t hat at t he
point of r epr oduct ion few if any r esour ces need be devot ed t o sur vival past r epr oduct ion.
16. Iteroparity (repeated reproduction)
(a) Or ganisms t hat pr oduce mor e t han one clut ch of offspr ing (pr ogeny) per life t ime ar e
said t o be it er opar ous (i.e., t o display it er opar it y). The advant age of it er opar it y is t hat
it allows or ganisms t o display mor e t han one st at ist ical “shot ” at pr oducing a successful
lit t er .
(b) “The cr it ica l fa ct or in t he evolut iona r y dilemma of big-ba ng ver sus r epea t ed
r epr oduct ion is t he sur vival r at e of t he offspr ing. If t heir chance of sur vival is poor or
inconsist ent , r epeat ed r epr oduct ion will be favor ed.”
Environmental Biology 321
POPULATION GROWTH
17 Population growth
(a) The simplest case of populat ion gr owt h is t hat which occur s when t her e exist no
limit at ions on gr owt h wit hin t he envir onment . In such sit uat ions t wo t hings occur
(i) The populat ion displays it s int r insic r at e of incr ease
(ii) The populat ion exper iences exponent ial gr owt h
18. Intrinsic rate of population increase (r
max
)
(a) The int r insic r at e of populat ion incr ease is t he r at e of gr owt h of a populat ion when
t hat populat ion is gr owing under ideal condit ions and wit hout limit s, i.e., as fast as it
possibly can. This r at e of gr owt h implies t hat t he differ ence bet ween t he bir t h r at e
and deat h r at e exper ienced by a populat ion is maximized.
(b) Not e t hat t he int r insic r at e of populat ion incr ease is a char act er ist ic of a populat ion
and not of it s envir onment . Indeed, in most envir onment s a populat ion is not able t o
achieve t his maximum r at e of gr owt h. However , a populat ion t hat is not gr owing
maximally can st ill exper ience exponent ial gr owt h.
(c) “A populat ion wit h a higher int r insic r at e of incr ease will gr ow fast er t han one wit h a
lower r at e of incr ease. The value of r
m ax
for a populat ion is influenced by life hist or y
feat ur es, such as age at t he beginning of r epr oduct ion, t he number of young pr oduced,
and how well t he young sur vive.”
19. Exponential growth
(a) Exponent ial gr owt h simply means t hat a populat ion’s size at a given t ime is equal t o
t he populat ion’s size at an ear lier t ime, t imes some gr eat er -t han-one number .
(b) For example, if a populat ion incr eased in size per unit t ime in t he following manner :
1, 2, 4, 8, 16, 32, 64, 128, et c. (or , e.g., 1, 3, 9, 27…, or 1, 5, 25, 125, …, et c. t hen t he
populat ion is displaying exponent ial gr owt h, each unit t ime t he populat ion is incr easing
by a fact or of 2 (or 3 or 5 in t he ot her examples; not e t hat exponent ial gr owt h is
occur r ing so long as t he r at e of incr ease per unit t ime is gr eat er t han a fact or of 1,
e.g., 2 or 4 or 10 or 1.2, et c.
(c) When populat ion size is gr aphed against t ime (e.g., gener at ions) a populat ion gr owing
exponent ially displays a J -shaped cur ve.
(d) Not e differ ences in int r insic r at es of gr owt h, in t his J -shaped cur ves, t hat r esult in
differ ences in r at es of exponent ial gr owt h (declining int r insic gr owt h r at es ar e seen
going fr om left t o r ight in t his gr aph):
(e) [In a r ich cult ur e medium bact er ia, gr own under aer obic condit ions, achieve a final
concent r at ion of 2-5 × 10
9
cells per ml in about 12-18 hour s. Alt hough plot t ed on a
differ ent t ime scale t he human gr owt h cur ve l ook s t h e s a me; t he human populat ion
at similar point s on t he gr owt h cur ve ar e shown.
(f) When populat ion size is gr aphed against t ime (e.g., gener at ions) a populat ion gr owing
exponent ially displays a st r aight line cur ve when gr aphed on semi-logar it hmic gr aph
paper (for example, below is a gr aph of t he exponent ial incr ease in t he comput er
322 CSIR-NET Life Sciences
pr ocessing power available per dollar —not e t hat on log-linear gr aph paper t his cur ve
is appr oximat ely a st r aight line):
(g) “The J -shaped cur ve of exponent ial gr owt h is char act er ist ic of populat ions t hat ar e
int r oduced int o a new or unfilled envir onment , or whose number s have been dr ast ically
r educed by a cat ast r ophic event and ar e r ebounding.”
(h) In ot her wor ds, a populat ion t hat is in an envir onment lacking limit s will gr ow
exponent ially (indeed, a populat ion t hat is capable of gr owing will t end t o gr ow
exponent ially), and t he r at e at which gr owt h will occur will be a funct ion of r
m ax
and
t he degr ee t o which t he envir onment mat ches t he ideal envir onment in which an
or ganism is capable of achieving r
m ax
.
20. Limits on population growth
(a) Exponent ial gr owt h cannot go on for ever ; sooner or lat er any populat ion will r un int o
limit s in t heir envir onment
21. Carrying capacity (K)
(a) “Populat ions subsist on a finit e amount of available r esour ces, and as t he populat ion
becomes mor e cr owded, each individual has access t o an incr easingly smaller shar e.
Ult imat ely, t her e is a limit t o t he number of individuals t hat can occupy a habit at .
100
80
60
40
20
0 20 40 60 80 100
time (generations)
number of individuals
B
i
l
l
i
o
n
s

o
f

H
u
m
a
n
s
6 Billion-1999
3 Billion-1960
N
u
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b
e
r

o
f

B
a
c
t
e
r
i
a
l

C
e
l
l
s
Time
Environmental Biology 323
Ecologist s define car r ying capacit y as t he maximum st able populat ion size t hat a
par t icular envir onment can suppor t over a r elat ively long per iod of t ime. Car r ying
capacity, sym bolized as K, is a pr oper t y of t he envir onment , and it var ies over space
and t ime wit h t he abundance of limit ing r esour ces.”
(b) In ot her wor ds, for any given or ganism, t her e will be a maximum number of individuals
t hat t he envir onment can suppor t wit hout t he envir onment being consequent ly
degr aded t o t he point wher e it can no longer suppor t t hat number of individuals.
(c) Gener ally, as populat ion size appr oaches car r ying capacit y, t he amount of some key
r esour ce declines per capit a t o t he point wher e individuals exper ience eit her a higher
deat h r at e or a lower fecundit y; t hus, as populat ion size appr oaches car r ying capacit y,
t he r at e of populat ion gr owt h declines t owar ds zer o.
22. Logistic growth
(a) Logist ic gr owt h is a mat hemat ical descr ipt ion of populat ion gr owt h t hat employs t wo
par amet er s, r
m ax
and K, and t wo var iables, N and t .
(b) The logist ic gr owt h cur ve is S-shaped.
(c) Se e F i gu r e : P op u l a t i on gr owt h p r e d i ct e d b y t h e l ogi s t i c mod e l .
(d) That is, t he populat ion gr ows exponent ially at a r at e which is det er mined by r
m ax
and
t he suit abilit y of a given envir onment t o an or ganism’s needs unt il populat ion size is
sufficient t hat t he limit at ions associat ed wit h t he car r ying capacit y of t he envir onment
ar e appr oached.
(e) This slows t he r at e of populat ion gr owt h in a way such t hat t he lar ger t he populat ion
becomes, t he slower it s r at e of gr owt h; t his slowing of t he gr owt h t r ansfor ms t he
cur ve fr om a J -shaped one t o an S-shaped one. Ult imat ely t he r at e of gr owt h of t he
populat ion r eaches zer o at t he car r ying capacit y.
(f) “Because t he r at e at which a populat ion gr ows changes wit h t he densit y of or ganisms
t hat ar e cur r ent ly in t he populat ion, t he logist ic model is said t o be densit y dependent .”
That is, populat ion gr owt h gr ows as populat ion densit y appr oaches t hat dict at ed by
an envir onment ’s car r y capacit y for t hat populat ion.
(g) Not e t hat populat ions do not t ypically display t he idealized logist ic gr owt h seen wit h
t he model. One deviat ion fr om idealized logist ic gr owt h is delayed feedback; t his can
cause populat ion size over shoot ing and, in fact , what is t ypically obser ved in r eal
100
80
60
40
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time
number of individuals
324 CSIR-NET Life Sciences
populat ions is not just effect s of r andom event s but also populat ions sizes which var y
up and down ar ound t he car r ying capacit y r at her t han r emaining invar iant exact ly at
t he car r ying capacit y.
23. K-selected populations (equilibrial populations)
(a) Idealized populat ions may be dist inguished in t er ms of t he logist ic gr owt h equat ion.
(b) For example, a species may bias it s life hist or y t owar d maximizing eit her r
m ax
or K
(c) That is, some or ganisms ar e good at incr easing t heir populat ion size r apidly in
envir onment s which lack limit s (e.g., weeds) while ot her species (e.g., gor illas) ar e
good at maint aining populat ion sizes at car r ying capacit y in envir onment s t hat have
limit s.
(d) A species t hat is bet t er at maint aining a populat ion at car r ying.
(e) capacit y in a st able envir onment is said t o be mor e K-select ed.
(f) A t ypical K-select ed species is shown t o t he r ight ®
24. r-selected populations (opportunistic populations)
(a) A species t hat is good at gr owing r apidly in, for example, dist ur bed envir onment s,
but is significant ly less capable of maint aining it s populat ion at car r ying capacit y in
undist ur bed (i.e., st able) envir onment s is t er med r-select ed.
25. r and K selection compared
(a) Few species ar e pur ely r- or K-select ed; e.g., t her e cer t ainly exist populat ions t hat
ar e able t o incr ease r apidly but may also t hr ive in mat ur e ecosyst ems. “It has been
difficult t o demonst r at e a dir ect r elat ionship bet ween populat ion gr owt h r at e and
specific life hist or y char act er ist ics. Incr easingly, ecologist s ar e r ecognizing t hat most
populat ions show a mix of t he t r adit ional r-select ed and K-select ed char act er ist ics;
life hist or y evolves in t he cont ext of a complex int er play of fact or s.”
Environmental Biology 325
(b) “Plant s and animals whose young ar e subject t o high mor t alit y r at es oft en pr oduce
lar ge number s of offspr ing. Thus, plant s t hat colonize dist ur bed envir onment s usually
pr oduce many small seeds, most of which will not r each a suit able envir onment .
Small size might act ually benefit such seeds if it enables t hem t o be car r ied long
dist ances… In ot her or ganisms, ext r a invest ment on t he par t of t he par ent gr eat ly
incr eases t he offspr ing’s change of sur vival.”
r K
Un st a ble en vi r on men t , St a ble en vir on men t , d en sit y
d en si t y i n d ep en d en t d ep en d en t i n t er a ct i on s
Or ganism size Sma ll Lar ge
Ener gy used t o make Low High
each individual
# Offspr ing pr oduced Many Few
Timing of mat ur at ion Ear ly Lat e
(wit h much par ent al car e)
Life expect ancy Short Long
Lifet ime r epr oduct ive event s One Mor e t han one
Sur vivor ship cur ve Type III Type I or II
POPULATION-LIMITING FACTORS
26. Density-dependent factors
(a) Densit y-dependent limit s on populat ion gr owt h ar e ones t hat st em fr om int r aspecific
compet it ion. Typically, t he or ganisms best suit ed t o compet e wit h anot her or ganism
ar e t hose fr om t he same species. Thus, t he act ions of conspecifics can ver y pr ecisely
ser ve t o limit t he envir onment (e.g., eat pr efer r ed food, obt ain pr efer r ed shelt er ,
et c).
(b) Act ions of t hat ser ve t o limit t he envir onment for conspecifics—e.g., eat ing, excr et ing
wast es, using up non-food r esour ces, t aking up space, defending t er r it or ies—ar e t hose
t hat det er mine car r ying capacit y. They ar e r efer r ed t o as densit y dependent because
t he gr eat er t he densit y of t he population, t he gr eat er t heir effect s. Densit y-dependent
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r-selected environmental instability
reduces population size before it
approaches carying capacity
number of individuals
K-selected-population
size is near carying capacity
due to density dependent factors
time (generations)
326 CSIR-NET Life Sciences
fact or s may exer t t heir effect by r educing bir t h r at es, incr easing deat h r at es, ext ending
gener at ion t imes, or by for cing t he migr at ion of conspecifics t o new r egions.
(c) “The impact of disease on a populat ion can be densit y dependent if t he t r ansmission
r at e of t he disease depends on a cer t ain level of cr owding in t he populat ion.” “A deat h
r at e t hat r ises as populat ion densit y r ises is said t o be densit y dependent , as is a bir t h
r at e t hat falls wit h r ising densit y. Densit y-dependent r at es ar e an example of negat ive
feedback. In cont r ast , a bir t h r at e or deat h r at e t hat does not change wit h populat ion
densit y is said t o be densit y independent … Negat ive feedback pr event s unlimit ed
populat ion gr owt h.”
(d) Pr eda t ion ca n a lso be densit y dependent since pr eda t or s oft en ca n swit ch pr ey
pr efer en ces t o ma t ch wh a t ever pr ey or ga n isms a r e most plen t ifu l in a given
envir onment . “Many pr edat or s, for example, exhibit swit ching behaviour : They begin
t o concent r at e on a par t icular ly common species of pr ey when it becomes ener get ically
efficient t o do so.
27. Density-independent factors
(a) Densit y-independent effect s on populat ion sizes (or st r uct ur es) occur t o t he same
ext ent r egar dless of populat ion size. These can be t hings like sudden changes in t he
weat her
(b) “Over t he long t er m, many populat ions r emain fair ly st able in size and ar e pr esumably
close t o a ca r r ying ca pa cit y t ha t is det er mined by densit y-dependent fa ct or s.
Super imposed on t his gener al st abilit y, however , ar e shor t -t er m fluct uat ions due t o
densit y-independent fact or s.”
28. Community
(a) A communit y consist s of all of t he or ganisms living wit hin a cer t ain geogr aphical
ar ea. These or ganisms include conspecifics as well as member s of ot her species.
These or ganisms int er act wit h each ot her bot h dir ect ly and indir ect ly.
(b) Numer ous (pessimist s might say “endless”) par amet er s affect what species ar e pr esent
and in what abundance
(c) “Simple gener a liza t ions ca n r a r ely expla in why cer t a in species commonly occur
t oget her in communit ies.”
(d) “The dist r ibut ions of most populat ions in communit ies ar e pr obably affect ed t o some
ext ent by bot h abiot ic gr adient s and int er act ions [wit h ot her species].”
29. Co-evolution
(a) Not only do t he abiot ic and biot ic component s of an ecosyst em impact on what species
ar e pr esent and in what abundance, but species also ar e modified by t heir int er act ions
wit h ot her species
(b) Co-evolut ion r epr esent s t he evolut ionar y modificat ion of or ganisms in r esponse t o
ot her or ganisms, par t icular ly when t wo or ganisms ar e mut ually modified in r esponse
t o modificat ions displayed by t he ot her , e.g.,
(i) a flower populat ion bet t er at t r act s cer t ain insect s which in t ur n evolve t o bet t er
exploit t he flower populat ion.
(ii) fast er r abbit s select for fast er coyot es which in t ur n select for fast er r abbit s.
Environmental Biology 327
COMMUNITY
COMMUNITY INTERACTIONS
1. Interspecific interactions
(a) Coevolut ion is one consequence of a mor e gener al cat egor y of ecology called int er specific
int er act ions (bet ween-species int er act ions). Pr eviously, we consider ed int r aspecific
int er act ions, i.e., t hose bet ween ver y similar or ganisms, conspecifics.
(b) Int er specific int er act ions r ange fr om t hose bet ween fair ly similar or ganisms t o t hose
bet ween ver y dissimilar or ganisms.
(c) A key dist inct ion bet ween int r aspecific and int er specific int er act ions is t hat t he for mer
but not t he lat t er shar e a gene pool;
(i) int r aspecific int er act ions do not gener ally lead t o t he ext inct ion of a species.
(ii) In int er specific int er act ions, loser s can go ext inct .
(d) Int er specific int er act ions include symbioses and can be cat egor ized as
(i) Pr edat ion/par asit ism (+/-)
(ii) Compet it ion (-/-)
(iii) Commensalism (+/0)
(iv) Mut ualism (+/+)
2. Commensalism
(a) Commensalism is a r elat ively unexploit ed int er specific int er act ion. The r eason for
t his has as much t o do wit h it s definit ion as anyt hing, i.e., commensalism is a
r elat ionship in which one member gains but t he ot her member neit her gains nor
loses; t his places commensalism on a knife’s edge bet ween pr edat ion and mut ualism.
(b) If t he “unaffect ed” individual is indeed affect ed, even just a lit t le, t hen t he r elat ionship
ca n no longer , t echnica lly, be t er med commensa lisms. I n t he r ea l wor ld, it is
essent ia lly impossible t o det er mine whet her t he “una ffect ed” member r ea lly is
unaffect ed, so t he concept is difficult t o apply. Never t heless, in absence of evidence
for mut ualism or pr edat ion t hen an assumpt ion of commensalisms is a r easonable
one.
3. Mutualism
(a) Mut ualisms, while not necessar ily as common as pr edat ion or int er specific compet it ion,
ar e st ill enor mously common. This makes some sense since a mut ualist ic r elat ionship
is one in which bot h member s gain. However , it is likely t hat most mut ualist ic
r elat ionships st ar t ed out , in evolut ionar y t ime, as exploit at ive (+/-) r elat ionships which
somehow wer e co-opt ed int o less exploit a t ive r ela t ionships. Exa mples include
ever yt hing fr om lichens, t o bees and flower s, t o mit ochondr ia and t he alr eady lect ur ed
on eucar yot ic cell.
PREDATION
4. Predation
(a) +/– int er act ions include
(i) Pr edat ion
(ii) Par asit ism
328 CSIR-NET Life Sciences
(iii) Par asit oidism
(iv) Her bivor y
(b) These int er act ions all involve
(i) one individual killing and t hen eat ing t he ot her fully (pr edat ion)
(ii) not killing and t hen eat ing t he ot her par t ially (par asit ism and her bivor y), or
(iii) let t ing one’s offspr ing do t he eat ing (par asit oidism).
(c) Not e t hat an addit ional kind of + – int er act ion does not involve eat ing but inst ead is
t he st ealing of some non-food a r esour ce fr om one individual by t he ot her : vines on
t r ees, for example, or a cow bir d’s br ood par asit ism.
5. Defense against predation
(a) Pr ey or ganisms display numer ous defenses against pr edat ion.
(b) That is, t her e exist s a number of defenses against + / – – (as well as + / –) int er act ions:
(i) Secondar y compounds (plant s)
(ii) Nut r it ional deficiencies (plant s)
(iii) Mechanical defenses (plant s)
(iv) Pr oduct ion of poisons (animals)
(v) Mechanical defenses (animals)
(vi) Running away & hiding (animals)
(vii) Fight ing back (most ly animals)
(viii) Cr ypt ic color at ion (most ly animals)
(ix) Bat esian mimicr y (animals)
(x) Müller ian mimicr y (animals)
(xi) Immune syst ems (animals)
(c) [in or der for a pr edat or t o obt ain benefit fr om pr ey t hey have t o encount er pr ey (i.e.,
be in close pr oximit y), t hen det ect pr ey (i.e., not ice t hat t hey cur r ent ly ar e in close
pr oximit y), t hen ca pt ur e pr ey, t hen successfully consume t he pr ey, a nd t hen
successfully der ive nut r ient benefit fr om t he pr ey, and minimally mor e benefit must
be der ived t han t he cost s of capt ur ing and consuming t he pr ey – hence, it is t o t he
pot ent ial pr ey’s benefit , as an individual or as a populat ion, t o minimize t heir number s
so as t o be r ar e and t her efor e r ar ely found by pr edat or s, t o be cr ypt ic in bot h color at ion
and behavior , t o be capable of escaping if not iced, t o be difficult t o consume or t o
digest , and t o not supply necessar y nut r ient s or t o be t oxin t o t he pr edat or in some
manner ].
6. Plant defenses against predation
(a) Of cour se, plant pr edat or s ar e called her bivor es.
(b) Typically a plant (and ot her st at ionar y or ganisms) will not manage t o achieve complet e
avoidance of pr edat ion, but inst ead will limit t heir own pr edat ion t o t hose or ganisms
t hat possess appr opr iat e mor phological or biochemical adapt at ions.
(c) It is impor t ant t o keep in mind t hat her bivor es can be big (cows) as well as small
(insect s, fungi, bact er ia) so mor e t han one defense is t ypically necessar y t o defeat all
possible pr edat or s.
Environmental Biology 329
(d) Of cour se, plant s also t end t o be eat en in pieces r at her t han as a whole or ganism, so
anyt hing a plant can do t o spar e par t of t he plant fr om being eat en can also be
advant ageous (t his r ule appar ent ly is also t r ue in t er ms of defenses against lawn
mower s).
(e) Plant defenses against pr edat ion include
(i) Secondar y compounds
(ii) Nut r it ional deficiencies
(iii) Mechanical defenses
7 Secondary compounds
(a) Secondar y compounds ar e chemicals t hat plant s pr oduce t hat ar e dist inct fr om t he
pr imar y met abolism t o some ext ent common t o all plant s. One r ole of secondar y
compounds ar e as defenses against pr edat ion, e.g., t oxins.
(b) What is t oxic t o one her bivor e may be useful t o anot her ; par t icular ly humans t ake
gr eat advant age of plant secondar y chemicals using t hem as dr ugs (bot h r ecr eat ional
and medicinal), spices, et c. Some animals (e.g., monar ch but t er flies) can act ually
incor por at e t hese t oxins int o t hemselves t o make t hemselves unpalat able t o some of
t heir own pr edat or s.
8 Nutritional deficiencies
(a) Plant s addit ionally t end t o lack cer t ain nut r ient s (e.g., essent ial amino acids).
(b) Such nut r it ional deficiencies for ce pr edat or s t o diver sify what plant s t hey consume,
t hus pr event ing her bivor es fr om get t ing t oo good (specialized) at exploit ing a par t icular
plant species.
9. Mechanical defenses
(a) Anyt hing a plant can do t o keep a her bivor e fr om r eaching, bit ing, or der iving benefit
fr om once a piece t hat has been r emoved can ser ve t o pr ot ect t he plant fr om being
consumed.
330 CSIR-NET Life Sciences
(b) Thor ns pr event lar ger t hings fr om comfor t ably eat ing a plant , while hair s and ot her
small appendages can keep small t hings fr om r eaching t he plant .
(c) Plant s also int er fer e wit h chewing by, essent ially, being less t han succulent , e.g.,.
t he shell of a nut or silica deposit ed in t he leaves of gr ass.
(d) (bet ween nut r it ional deficiencies, mechanical defenses, and secondar y compounds
one speaks of low for age qualit y and it is plant s t hat r epr esent low-qualit y for age t hat
t end t o accumulat e when her bivor e pr essur es ar e high, i.e., high animal t o plant
r at ios).
10. Animal defenses against predation
(a) Animals ar e a lit t le bit mor e ver sat ile behavior ally when it comes t o defending
t hemselves against pr edat ion.
(b) For example, animals can
(i) Run and hide (par t icular ly t he lat t er aided by cr ypt ic color at ion)
(ii) Pr oduce poisons t hat make t hem unpalat able
(iii) Employ mor phological adapt at ions t hat int er fer e wit h consumpt ion
(iv) Fight back using bot h mor phological and chemical defenses (some plant s, t oo,
can fight back using, for example, act ively spr ayed chemical defenses)
(v) Not looking like pr ey (cr ypt ic color at ion)
11. Cryptic coloration
(a) Cr ypt ic color at ion is camouflage, t he ar t of looking like somet hing else, i.e., not
hiding behind somet hing but inst ead not being visible against appr opr iat e backgr ounds.
(b) [“Camouflage, called cr ypt ic color at ion, is t he quint essent ial passive defense, making
pot ent ial pr ey difficult t o spot against it s backgr ound. A camouflaged animal need
only r emain st ill on an appr opr iat e subst r at e t o avoid det ect ion”].
12. Aposematic coloration
(a) A differ ent appr oach is t o t ast e bad (or be unpalat able for var ious ot her r easons) and
t o adver t ise t his
(b) Aposemat ic color at ion is how or ganisms adver t ise unpalat ableness, at least visually
(humans, or cour se, display a dist inct bias in t er ms of sensor y input hence we t end t o
not ice t he visual displays by animals much mor e so t han, for example, t he olfact ic
displays – not ice t hat since bir ds display t he same bias we oft en int er pr et t hese visual
displays in t er ms st r at egies t hat int er fer e wit h pr edat ion by bir ds).
(c) For example, t he black and yellow st r ipes on bees r epr esent aposemat ic color at ion,
and it wor ks!
(d) Aposemat ic color at ion is so successful, in fact , t hat it gives r ise t o mimicr y
(i) Bat esian mimicr y
(ii) Müller ian mimicr y
13. Batesian mimicry
(a) Bat esian mimicr y is t he t endency of palat able and ot her wise succulent pr ey species
t o pr et end t o be unpalat able by looking like unpalat able species. This wor ks t o a
Environmental Biology 331
point , but limit s t he size of t he mimic’s populat ion since once mimics ar e sufficient ly
pr evalent , pr edat or s will cat ch on t o t he mimicr y.
(b) However , when t heir number s ar e sufficient ly few, t he mimic gains fr om pr ot ect ion
fr om pr edat ion while simult aneously not put t ing out t he r esour ces needed t o achieve
lack of palat abilit y, et c.
14. Müllerian mimicry
(a) Mimicr y wor ks in ever ybody’s favor when unpalat able species mimic each ot her (e.g.,
bot h wasps and bees shar ing black and yellow aposemat ic color at ion). Such mimicr y
incr eases t he r epr esent at ion of lack of palat ableness among pot ent ial pr ey associat ed
wit h a given for m of aposemat ic color at ion.
(b) Not e t hat bot h Bat esian and Müller ian mimicr y can occur simult aneously wit h t he
same aposemat ic color at ion wit hin t he same communit ies (i.e., a gr oup a similar ly
mar ked or ganisms, some of which ar e har mless and ot her s which ar e not ).
15. Predation and species diversity (keystone species)
(a) Anot her way t hat t wo dir ect ly compet ing species can achieve coexist ence r esult s
fr om pr edat ion. A pr edat or t ypically feeds on mor e t han one species. By doing so,
t hey ser ve t o keep t he populat ions of bot h species below t he sizes one or bot h could
at t ain in t he absence of pr edat ion (i.e., below car r ying capacit y). This can allow bot h
compet ing species (i.e., t he pr ey) t o coexist , especially if t he weaker compet ing species
happens t o be bet t er at escaping pr edat ion.
(b) Addit ionally, opt imal for aging can r esult in pr ey caught and consumed as a funct ion
of t heir populat ion densit ies such t hat pr edat ion maint ains pr ey diver sit y by fr equency-
dependent effect s in t he same manner t hat fr equency-dependent select ion can maint ain
a balanced polymor phism.
(c) One can descr ibe a pr edat or whose pr esence has a pr ofound impact on t he species
diver sit y of a given communit y as a keyst one species.
COMPETITION BETWEEN SPECIES
16. Interspecific competition
(a) Int er specific compet it ion r epr esent s a lose-lose int er act ion (–/–), t hat is, bot h species
ar e less able t o conver t r esour ces int o pr ogeny because t he ot her species is laying
claim t o t he same r esour ces.
(b) Not e t hat t his is an unst able sit uat ion t hat will t end t o select for eit her bet t er means
of acquir ing t he cont est ed r esour ces, or a swit ching t o a differ ent r esour ce.
(c) Addit ionally, not e t hat while a compet ing species may be mor e effect ive in exploit ing
any given r esour ce, conspecifics will always be compet ing wit h any given individual
for a lar ger var iet y of r esour ces t han will int er specifics.
(d) Thus, gr owt h of a given species may be limit ed by bot h conspecifics (int r aspecific
compet it ion/densit y-dependent fa ct or s) a nd int er specific compet it ion (a densit y-
independent fact or ).
332 CSIR-NET Life Sciences
(e) “Expr ession” of t he cost s of int er specfic compet it ion include:
(i) Compet it ive exclusion
(ii) Resour ce par t it ioning
(iii) Char act er displacement
(iv) Fundament al vs. Realized Ecological Niche
17. Interference competition
Int er specific compet it ion t hat involves act ual int er specific fight ing is t er med int er fer ence
compet it ion.
18. Exploitative competition
Int er specific compet it ion which involves no fight ing but inst ead a co-usage of one or mor e
r esour ces is t er med exploit at ive compet it ion.
19. Competitive exclusion
(a) “Two species wit h similar r equir ement s cannot coexist in t he same communit y; one
species would inevit ably har vest r esour ces and r epr oduce mor e efficient ly, dr iving
t he ot her t o local ext inct ion. Even a slight r epr oduct ive advant age would event ually
lead t o t he eliminat ion of t he infer ior compet it or and an incr ease in t he densit y of t he
super ior one”. This is t he compet it ive exclusion pr inciple.
(b) Two populat ions wit h ver y similar needs, living sympat r ically, will be in t oo gr eat a
compet it ion wit h each ot her t o coexist , unless bot h populat ions ar e t he same species
(in which case, of cour se, t hey wouldn’t be t wo separ at e populat ions). Similar ly, t wo
populat ions can coexist if t heir needs sufficient ly differ .
20. Resource partitioning
(a) Ext inct ion of one of t wo populat ions living sympat r ically and compet ing over t oo
many r esour ces is not t he only possible out come of int er specific compet it ion. An
a lt er na t ive out come is t he evolut ion of a diver gence of r esour ce needs. Such a
diver gence is called r esour ce par t it ioning, and is simply t he ecological ver sion of t he
idea t hat it is oft en easier t o swit ch t han it is t o fight .
21. Character displacement
(a) Char act er displacement is pr esumably a consequence r esour ce par t it ioning. “The
t endency for char act er s t o be mor e diver gent in sympat r ic populat ions of t wo species
t han allopat r ic populat ion of t he same t wo species is called char act er displacement .”
(b) That is, char act er s diver ge pr esumably in r esponse t o int er specific compet it ion, but
do not diver ge in populat ions not subject t o t he same int er specific compet it ion.
(c) Th u s , t h e ch a r a ct er differ s bet ween t h e popu la t ion u n der goin g in t er s pecific
compet it ion and t he populat ion not under going int er specific compet it ion.
22. Ecological niche
(a) What is being fought over in int er specific compet it ion is var ious aspect s of t he ecological
niche. A niche is t he sum t ot al of what an or ganism does in it s envir onment , including
all of t he r esour ces consumed.
Environmental Biology 333
23. Fundamental niche
(a) All of t he r esour ces a populat ion could exploit under ideal condit ions, wher e t her e
exist s no int er specific compet it ion, is t er med t he fundament al niche of an or ganism.
The fundament al niche basically r epr esent s as good as t hings can get for an or ganism.
A populat ion able t o exploit it s fundament al niche would be able t o achieve it s maximal
populat ion size.
24. Realized niche
(a) Not hing, of cour se, is per fect , and t he fundament al niche r epr esent s per fect ion t o
t he exploit ing populat ion. In t he r eal wor ld, populat ions do not have access t o all of
t he r esour ces t hey could possibly exploit . Such a limit at ion on r esour ce acquisit ion is
t er med a r ealized niche, i.e., what r esour ces a populat ion can exploit in a r eal
envir onment , par t icular ly one in which int er specific compet it ion occur s.
TROPHIC STRUCTURE
25. Trophic structure
Tr ophic st r uct ur es ar e t he feeding r elat ionships wit hin communit ies and t her efor e wit hin
ecosyst ems, t hat is, who’s eat ing whom.
26. Trophic level
Tr ophic levels r efer t o how far r emoved fr om t he or iginal sour ce of ener gy an or ganism is
wit hin a t r ophic st r uct ur e.
27. Primary producer
The fir st t r ophic level is made up of t he pr imar y pr oducer s, t he or ganisms t hat obt ain fr om
inor ga nic sour ces t he ener gy t ha t power s ecosyst ems. Pr ima r y pr oducer s t ypica lly a r e
phot osynt het ic or ganisms. Mor e gener ally, pr imar y pr oducer s ar e aut ot r ophs (i.e., t hey fix
CO
2
).
28. Consumers
Consumer s ar e t he het er ot r ophs, i.e., or ganisms t hat obt ain t heir car bon fr om ot her or ganisms.
The t ypical consumer is a chemohet er ot r oph t hat consumes ot her or ganisms or par t s of ot her
or ganisms t o obt ain t heir car bon and ener gy. In addit ion t o t he t ypes of consumer s list ed below
we can also speak of omnivor es, i.e., consumer s t hat eat at differ ent t r ophic levels including
consuming pr oducer s and det r ivor es, which ar e consumer s t hat consume det r it us which is t he
br oken up r emains of or ganisms].
29. Primary consumer (herbivore)
A pr imar y consumer is a consumer t hat eat s pr imar y pr oducer s. Pr imar y consumer s ar e called
h e r b i vor e s .
30. Secondary consumers (carnivore)
Secondar y consumer s eat pr imar y consumer s.
334 CSIR-NET Life Sciences
31. Tertiary consumers
Ter t iar y consumer s eat secondar y consumer s.
32. Decomposers
(a) Decomposer s consume t he wast e given off by living or ganisms or t he r emains of dead
or ganisms which t hey did not kill.
(b) “The or ga nic ma t er ia l t ha t composes t he living or ga nisms in a n ecosyst em is
event ually r ecycled, br oken down and r et ur ned t o t he abiot ic envir onment in for ms
t hat can be used by plant s. Decomposer s, which feed on nonliving or ganic mat er ial,
ar e key t o t his r ecycling pr ocess. The most impor t ant decomposer s ar e bact er ia and
fungi, which fir st secr et e enzymes t hat digest or ganic mat er ial and t hen absor b t he
br eakdown pr oduct s; some can even digest cellulose.”
(c) “In fact , all het er ot r ophs, including humans, ar e decomposer s in t he sense t hat t hey
br eak down or ganic mat er ial and r elease inor ganic pr oduct s, such as car bon dioxide
and ammonia, t o t he envir onment .”
33. Food chain
A simplificat ion of t he t r ophic st r uct ur e of an ecosyst em is t he food chain. Food chains r efer t o
t he passage of nut r ient s and ener gy fr om a pr imar y pr oducer t o a pr imar y consumer t o a
secondar y consumer , and soon.
34. Food web
(a) Far mor e r ealist ic is t he concept of food webs. Food webs ar e like food chains but
mor e r ealist ic, i.e., allowing for species t o consume mor e t han one ot her kind of
species. In addit ion, food webs allow individual species t o consume at mor e t han one
t r ophic level.
(b) For exa mple, huma ns consume pr ima r y pr oducer s (e.g., soya bea ns), pr ima r y
consumer s (e.g., cows), and secondar y (or higher ) consumer s (e.g., salmon).
(c) [grazing food chains have pr oducer s at base which t he her bivor es t hen gr aze on.
While gr azing food chains ar e impor t ant , in nat ur e t hey ar e out number ed by det rit us-
based food chains. In det r it us-based food chains, decomposer s ar e at t he base of t he
food chain, and sust ain t he car nivor es which feed on t hem. In t er ms of t he weight (or
biomass) of animals in many ecosyst ems, mor e of t heir body mass can be t r aced back
t o det r it us t han t o living pr oducer s.
ECOLOGICAL SUCCESSION
35. Ecological succession
(a) One t hing t hat limit s t he car r ying capacit y for many or ganisms is t hat t he pr esence
of t hese or ganisms essent ially spoils t he envir onment for t heir cont inued pr esence.
Su ch or ga n isms t ypica lly a r e r -s e l e c t e d , a nd essent ia lly a r e good a t finding
envir onment s t hey can exploit , exploit ing t hose envir onment s, t hen giving way t o
or ganisms which ar e bet t er at hanging on in t hose envir onment s.
Environmental Biology 335
(b) The exploit at ion of an envir onment by one populat ion, followed by t he exploit at ion by
a second (t hir d, et c.) populat ion is t er med ecological succession.
(c) “Many of t he changes in communit y st r uct ur e dur ing succession may be induced by
t he or ganisms t hemselves. Dir ect biot ic int er act ions may be involved, including
inhibition of some species by ot her s t hr ough exploit at ive compet it ion, int er fer ence
compet it ion, or bot h. The pr esence of or ganisms also affect s t he abiot ic envir onment
by modifying local condit ions. This may r esult in facilitation, in which t he gr oup of
or ganisms r epr esent ing one st age ‘paves t he way’ for species t ypical of t he next st age
. . . Somet imes t he changes t hat facilit at e t he development of a lat er st age act ually
make t he envir onment s unsuit able for t he ver y species r esponsible for t he changes.”
(d) Ecological succession continues in a habitat until species, typically K-s e le ct e d , t hat
ar e good at nur t ur ing t heir young wit hin t he same envir onment (as well as good at
excluding ot her species) comes t o dominat e t he envir onment , or unt il cat ast r ophic
change essent ially wipes t he slat e clean, making an envir onment once again exploit able
t o t he r-select ed populat ions.
36. Primary succession
(a) Ecological succession t ypically occur s in fair ly well-defined waves of succeeding
or ganisms. When t he envir onment being exploit ed is essent ially lifeless—lacking in
bot h living or ganisms and in t heir r emains—t hen t he fir st r ound of exploit at ion is
t er med pr imar y succession.
(b) Pr imar y succession occur s, for example, following volcanic or glacial dest r uct ion of
an envir onment . The fir st or ganisms t hat exploit an ot her wise lifeless t er r ain ar e
t er med pr imar y successor s. This is a fair ly r ar e occur r ence especially r elat ive t o t he
much-mor e familiar secondar y succession t hat we obser ve in dist ur bed habit at s all
ar ound us.
37. Secondary succession
(a) Seconda r y succession is succession t ha t follows pr ima r y succession, i.e., of a n
envir onment t hat alr eady cont ains life (or , at least , soil).
(b) “Because r esour ce availabilit y changes over t he cour se of succession, differ ent species
compet e bet t er at differ ent st ages. Ear ly st ages ar e t ypically char act er ized by r-select ed
species t hat ar e good colonizer s because of t heir high fecundit y and excellent disper sal
mechanisms. Many of t hese may be descr ibed as ‘fugit ive’ or ‘weedy’ species t hat do
not compet e well in est ablished communit ies, but maint ain t hemselves by const ant ly
colonizing newly dist ur bed ar eas befor e bet t er compet it or s can become est ablished in
t he same places.”
38. Climax community
(a) The communit y wit hin an ecosyst em t hat exist s following ecological succession is
t er med t he climax communit y. A climax communit y is made up of or ganisms t hat ar e
good at r epr oducing in t he face of int er specific compet it ion
(b) “At t he climax st age, envir onment al condit ions ar e such t hat t he same species can
cont inue t o maint ain t hemselves. For example, t he [maple-beech] for est t hat is t he
climax st age of old-field succession [in much of Ohio] maint ains t he moist , shaded
336 CSIR-NET Life Sciences
envir onment t hat allows offspr ing of t hese species t o gr ow, while inhibit ing most of
t he species t ypical of ear lier st ages of succession.”
(c) Climax communit ies will r emain in place unt il eit her t he climat e changes, a bet t er
compet it or ar r ives, or t he communit y is cat ast r ophically disr upt ed, e.g., by fir e or ,
mor e r ecent ly, by ext ensive logging;
ISLAND BIOGEOGRAPHY
1. Island biogeography
(a) In or der for an ecosyst em t o go t hr ough succession, t he or ganisms in each wave of
succession must be available in t he local envir onment . The far t her an ecosyst em is
fr om a sour ce of t hese or ganisms, t he less likely t hese or ganisms will be pr esent and
t her efor e t hat succession will occur . The smaller an island is, t he less likely t hat
species will find t heir way t o t he island and t he mor e likely t hat species pr esent on
t he island will go ext inct (due t o smaller size and due t o r esult ant ly smaller populat ions,
r espect ively).
(b) This can be seen most obviously on islands: t he far t her an island is fr om a sour ce of
or ganisms, t he less likely t he given or ganisms will find t heir way t o t he island. The
flip side is t hat as a consequence of, if not hing else, r andom ext inct ion, t he smaller
an ecosyst em is, t he less able it is t o hold on t o t he species t hat it has. Thus, t he
far t her an island or ecosyst em is fr om ot her islands or ecosyst ems, and t he smaller
t he island or ecosyst em, t he mor e impover ished of species eit her is likely t o be.
(c) Applicat ion of t hese ideas t o our envir onment is somewhat pr ofound because t hey
t ell us t hat we can’t go on dest r oying ecosyst ems for ever wit hout r isking t heir ver y
exist ence. In ot her wor ds, event ually if we conver t ever y last for est int o far mland,
housing t r act , or par king lot , t he r emnant s of ecosyst ems will be so small t hat t hey
will be unable t o sust ain what species t hey st ar t wit h, and ecosyst ems will be so far
apar t t hat t hey will be unable t o r eacquir e species fr om similar ecosyst ems
(d) This essent ially, ult imat ely r epr esent s a genet ic bot t lenecking of t he ent ir e wor ld,
and if t he goal of humans is t o sur vive past t his envir onment al disast er of our own
making, t hen t he big loser s will most definit ely be our selves.
FLOW OF ENERGY THROUGH ECOSYSTEMS
1. Ecosystem
(a) “An ecosyst em consist s of all t he or ganisms living in a communit y as well as all t he
abiot ic fact or s wit h which t hey int er act .”
(b) Not e t hat t he boundar ies of ecosyst ems ar e t ypically not ar bit r ar ily defined, but inst ead
ar e defined in some meaningful way: A pond, a field, a for est , et c.
(c) Ecosyst ems ar e t ypically under st ood in t er ms of
(i) Ener gy flow t hr ough ecosyst ems
(ii) Chemical cycling wit hin (and t hr ough) ecosyst ems
Environmental Biology 337
(d) Not e t hat bot h involve t he movement of “st uff” t hr ough bot h biot ic and abiot ic
component s of t he ecosyst em.
(e) “Ecosyst ems ecologist s view ecosyst ems as ener gy machines and mat t er pr ocessor s.
By gr ouping t he species in a communit y int o t r ophic levels of feeding r elat ionships,
we can follow t he t r ansfor mat ion of ener gy in t he whole ecosyst em and map t he
movement s of chemical element s as t hey ar e used by t he biot ic communit y.”
2. Energy flow
(a) Ener gy does not cycle t hr ough ecosyst ems but inst ead ent er s ecosyst ems and is used
up wit hin ecosyst ems. Ult imat ely, ener gy is lost fr om ecosyst ems pr imar ily as wast e
heat , t he most t her modynamically unavailable for m of ener gy.
(b) “Ener gy ent er s most ecosyst ems in t he for m of sunlight . It is t hen conver t ed t o
chemical ener gy by aut ot r ophic or ganisms, passed t o het er ot r ophs in t he or ganic
compounds of food, and dissipat ed in t he for m of heat . . . The movement s of ener gy
and mat t er t hr ough ecosyst ems ar e r elat ed because bot h occur by t he t r ansfer of
subst ances t hr ough feeding r elat ionships. However , because ener gy, unlike mat t er ,
cannot be r ecycled, an ecosyst em must be power ed by a cont inuous influx of new
ener gy fr om an ext er nal sour ce (t he sun). Thus, ener gy flows t hr ough ecosyst ems,
while mat t er cycles wit hin t hem.”
Sun
Plants
Animals
Soil
Organic
Matter
Space
Environment
Plants
Animals
Soil
Organic
Matter
Solar
Radiation
Dispersal
Migration
Erosion
and
Deposition
Longwave
Radiation
Heat
Dispersal
Migration
Erosion
and
Leaching
Consumers
Decomposers
Plants
Ecosystem
Photosynthesizing
Organisms
Herbivores
Carnivores
“Pyramid of Life”
Photons from Sun
338 CSIR-NET Life Sciences
(c) Not e t hat ener gy flows t hr ough ecosyst ems most ly as bonds bet ween car bon at oms
and bonds bet ween car bon and hydr ogen at oms, e.g., as one finds in car bohydr at es
and lipids; consequent ly, wit hin and bet ween or ganisms t he car bon cycle and t he flow
of ener gy ar e quit e similar , at least unt il t he t wo ar e decoupled in t he cour se of
cellular r espir at ion (i.e., t he separ at ion of car bon at oms fr om t heir ener gy).
PRODUCTIVITY
3. Primary productivity
Only a small fr act ion of t he sunlight st r iking t he ear t h is conver t ed t o chemical ener gy by
pr imar y pr oducer s. That sunlight ener gy t hat is conver t ed t o chemical ener gy, over a given
per iod, is t er med pr imar y pr oduct ivit y.
4. Gross primary productivity
Gr oss pr imar y pr oduct ivit y is all of t he light ener gy t hat is conver t ed t o chemical ener gy by
pr oducer s.
5. Net primary productivity
Net pr imar y pr oduct ivit y is all of t he light ener gy t hat is conver t ed t o chemical ener gy and t hat
is subsequent ly st or ed by t he pr imar y pr oducer (i.e., t he gr oss pr imar y pr oduct ivit y minus t hat
employed t o r un t he pr imar y pr oducer ’s met abolism). The r at io of net pr imar y pr oduct ivit y t o
gr oss pr imar y pr oduct ivit y gives an indicat ion of t he cost of keeping t he or ganism going, wit h
lar ge r at ios indicat ive of r elat ively few cost s (e.g., algae, ~50%) and smaller r at ios associat ed
wit h many cost s (e.g., complex plant s such as t r ees, ~10%).
6. Biomass
Net pr imar y pr oduct ivit y is st or ed as biomass (dr y mass of or ganisms).
7. Standing crop biomass
St anding cr op biom