Plant nutrition

Nutrition is the process of acquiring energy and material s, while energy can be simply
defined.as capac ity to do thi1.igs. Energy can nei ther be created nor destroyed, but it occurs in
various forms and can be convened from one form to another. The co.mmon uses of energy
include but not limited to the following: e lectri cal transmi ssion of nerve impul ses, generation~
of heat during respirati_on to mai ntain hody temperature in birds and mammals,
biolumniscen ce-productio n of light by fireflies and glow worms, mechanical contraction of
muscles, synthesis of new substances for growth and repair, and active transport of materials
in and out of the cells. Carbon, is the most fundamental materials needed by living organisms.
OjhCJ"S include 02, CO2, and macro and micro elements. Thus, living organisms are grouped
on the basis of sources of energy and carbon. Though several sources of energy exist, only
two: Light and chemical energy are suitable sources for use by living organisms. Autotroph
(autos, self>, is an organism capable of synthesizing its own organic substances from
inorganic .compounds. Autotrophs produce their own sugars, lipids, and amino acids using
carbon dioxide as a source of carbon, and ammonia or nitrates as a source of nitrogen.
Organisms that use light for the energy to synthesize organic compounds are called
photosynthetic autotrophs; organisms that oxidize such compounds as hydrogen s~
(H2S) to obtain energy are called chemosynthetic autotrophs, or chemotrophs.
Photosynthet ic autotrophs include the green plants, certain algae, and the pigmented sulfur
bacteria. Chemotroph s include the iron bacteria, the nitrifying bacteria, and the non-
pigmented sulfur bacteria. Heterotrophs (heteros, others) are organisms that must obtain
their energy from organic compounds (all forms of sugar as sources of carbon). While
autotrophs are the producers in the ecosystem, heterotrophs , which depend on their (autotroph)
products are either consumers, detritivores. or decomposers. .
.

Table 2.1: Nuritional categories of organisms

Carbon/energy Autotrophic (inorganic carbon) Heterotrophic (o~anic

source carbon)
Phototrophic (light Photoautotro phic, eg blue green algae. Photohetero trophic eg purp!~
energy) (Anabaena, Spirulina), green algae and non-sulphur bacteria
plants
Chemotrophic Chemoautot rophic eg, Nitrosomon.as Chemoheter otrophic eg.
(Chemical energy) and Nitrobacter saprophytes , most bacteria,
•
parasites and animals

Photosynthesis '• '!.

A •
Photosynthe sis is the process by which plants, some bacteria, and some protistans u~
fhe energy from sunlight to pr~uce sugar, which cellu~_~ respiration converts into ATP, the / ·
"fuel" used by aH Jiving things. The conversion of unusable· sunlight energy into usabl~: '
I

chemical energy, is associated with the actions of the green pigment chlorophyll. Most of the
~:ti)ne, the phot9 s ynthetic process. uses water and releases the oxygen that we absolutely ft\PSl
47 .
 ha ve to sta y alive, anJ we need the food as well. The ove rall reacti on o f thi s process can he
written as:

The ahove chemica l equati on translates as, in the presence of c hlorophyll and sunli ght, six
molec ules of water plus six mnlcc ulcs of carbon di ox ide produce one mo lecule of sugar plus
six mo lecules o f oxygen. This equation for photosynthesis is a deceptively simple summary of
a complex process. In rea lity, photosynthesis is not just one reaction but two processes each
with multiple steps. To produce a sugar molecule such as sucrose, plants require nearly 30
distinct protei ns that work wi thin a complicated membrane structure. Research into the
mechanism of photosynthesis centers on understanding the structure of the photosynthetic
components and the molecular processes that use radiant energy to drive carbohydrate
synthesis. The researc h invol ves several disciplines, including physics, biophysics, chemistry,
structural biology, biochemistry, molecular bi o logy and physiology, and serves as an
outstandjng example of the success of multidisciplinary research. As such, photosynthesis
presents a spec ial challenge in understanding several interrelated molecular processes.
Photosynthesis is not a very efficient process, however it is the source of food (carbon anu
energy). and oxygen on earth for life . Of the sunlight reaching the surface of a leaf.
approximate! y:
~~"
,--' \
I . ~ ~
('\ .• 75% is evaporated
15% is reflected
~L~ • 5% is transmitted through the le
~
~ 9) • 4% is converted to heat energy
• 1% is used in photosynthesis ~v,0 ,

The two stages of photosynthesis are known as the !igK reactions (photo part) and the
dark reactions (the synthesis part, also known as the Calvin cycle). These two stages of
convertin ~ carbon dioxide and water into glucose are almost the· same for all plants, algae and
photosy 1thesizing bacteria. In the following section, both stages will be described and
hopefr ly it will become clear how the food we eat can be created from the light of the sun.
PhotosjTithesis is carried out by many different organisms, ranging from plants to bacteria.
The best known form of photosynthesis is the one carried out by higher plants and algae,~
well as by cyanobacteria and their r~latives, which are responsible for a major part of
photosynthesis in oceans. About 40% of photosynthesis is carried out by algat
(phytoplankton), the maj~r source of food in the aquatic environment. All thes~ organisin'
convert CO 2 (carbon dioxi~e) to organic material_ by redu~ing this. gas to _carbohydrates .i:; (
complex set of reactions. ~e~gos (Qr th1S..f$UJJCllon reaction ult1matel y c o ~
.... r throu gh a s~ e n . ~ reactions calkiLQ!t~ly.§~: which is then converted to oxyg~
a1!?, protons. Energy rior t h"is process i_
·s provided by light, · rnell"'
which is absorbed by ~ig ~
(primarily chlorophylls . and ~arotcno1ds) . . C:hloro~hylls absorb blue and red -~
ca~ote_!l_~?~ blue-~~!!_g_~t, but green_~~ YC?~~~ li_ght ~_e_J1ot~f~_tj_~ely _abs_5>~eS(
-- ______
photosynthetic_p
...
•~~ ~-_e!!n~ th erefore, hght of these colors is either reflected by 1
or passes through the leaves. This is why plants are green
. Oth~r pho~osynthetic organ isms,
such as cyano bacte ria (form erly kn~~~ as_b~ue-ire~n algae
) and re~ _~lgae, have . a4ditional
pigments called _£h ycobi \ins that are red or blue and that absor
b the colors of visible light that
are not effectively absor bed by chlor ophy ll and carotenoids.
Yet other organisms, such as the
purple and green bacte ria (whic h, by the way, look fairly
brown under many growth
conditions), conta in ~acte rioch lorop hyll l hat absorbs in the
infrared, in addition to in the blue
part of the spect rum. Thes e bacte ria d~ not evolv e oxyg en,
but perform photosynthesis under
anaerobic (oxyg en-le ss) cond itions . Thes e bacteria effici
ently use infrared light for
photosynthesis. Infra red is light with wave lengt hs above 700
nm that cannot be seen by the
human eye~ some bacte rial speci es can use in_frared light with
wavelengths of up to 1000 nm.
However, most pigm ents are not very effective in absor
bing ultraviolet light (<400 nm),
which also canno t be seen by the huma n eye. Light with wave
lengths below 330 nm becomes
increasingly dama ging to cells, but virtually all light at these
short wavelengths is filtered out
by the atmo spher e (mos t prom inent ly the ozon e layer) befor
e reaching the earth. Thou gh most
plants are capab le of produ cing comp ound s that absor
b ultraviolet light, an increased
exposure to light aroun d 300 nm has detrimental effects on plant
productivity.

--, , ~ '•
~ t ",: > ✓ · :.

,
2. 1·.. o·1agram of a typical plant, showing the inputs and outputs of the photosynthetic
.
F1g.
process

· . f hotosynthetic organisms: leaves from higher plants flanked

Fig 2 2· Exam p1es O P by
· · · . · h · · h)
ynthetic purple bacteria (left) an<;i cyanobactena
r.olom es of P o (ng t •
 waH·knr th 111111 )

Fig. 2.3: Absorption spectrum of isolated chlorophyll and carotenoid species. The color
associated with the various wavelengths is indicated above the graph.

Fig. 2.4 : A typical 'plant cell

Light plays a very important role as the driving force in the process of photosynthesis.
Knowing this , we can observe where photosynthesis actually occurs in a photosynthetic
organism . The many structures present in th~ cell cytoplasm (the entire contents· of the cell,
not incl~1din g the nucleus. and sunounded by the plasma membrane) are organelles, which
you can think of as "little organs." each having a specialized function of its own. ~
~ type of_J))cmi)ra t ~und organelle , the most !me_ortant of wh~js the chloropl~ e site
~ of photosynth L·S1'.-. 111 plants. Chloroplasts carry out photosynthesis, c onverting sunlight to
chemical ene rgy 's tored in sugar and other organic molecules. An eas y way to remember this
11
1s to think ul chloroplasts as chemical factories powered by the sU ·_
01
,\ 11 green part s or a plant contain chloroplasts. The \eaves, however, are the major organs
11 1illwsynthew , in most plants. I). leaf is comprised of a waxy u·p per and lower layer,_E-~
' 11 11J_illc_lH) Cr (th1.: mcsoph yll ) in between , kind of like a sandwich. This middle layer is the
.---
50
portion that is specialized for photosynthesis . Actually, there are two parts to this middle layer:
there are cells that are long and columnar in shape on top, and there are spongy cells with lo ts
of air spaces right below• The top layer filters light to the spongy layer, where photosynthes is
occurs. Other important structures on a leaf (or more specifically, the waxy layers) are the
stomata, tiny pores with specialized cells called &ard cells on each side. Each' stoma is
~ aii; a gap between a pair of guard cells. In addition, there~are also specialized cells for the
transport of nutrients and water. Within a leaf, veins subdivide repeatedly and branch
throughout the middle layer of the leaf. Veins consist of water-conducti ng tissue 0!1 top of
nutrient-conduc ting tissue, surrounded by a sheath. Hence the term, bundle-sheath ceUs. The
location of these veins brings the conducting tissues in close contact with the photosynthetic
tissue, which obtains water and minerals from the water-conducti ng tissue and loads it sugars
and other organic products into the ir'utrient-condu cting tissue for shipment to other parts of
the plant.
Chloroplasts, found mainly in the middle layer of the leaf, are bound by a double
_ h: Cl 1.,vt r x. v\
membrane that encloses the stroma, tfle dense fluid content of the chloroplast. Another
important structure to become familiar with is a thylakoidJ0i.icbis a flattened membran~ _ac
inside the chlor~~sts,~ sed to convert light energy to chemic!ll ~~Y· Membranes of the
thylakoid system separate the stroma from the thylakoid space. Thylakoids are concentrated in
,c,-l
C)J ~ r- )(."' •
stacks ca1led grafia. Thylakoids can be pictured as stacks of pancakes, immersed m heavy
maple syrup (which you can think of as the stroma).

Fig. 2.S: The anatomical structure of a leaf.

 Fig. 2.6: The thylakoids

Pigment types
As earlier mentioned, leaves are the major sites of photosynthes is. What makes a leaf
green is chlorophyll , the green pigment located within the chloroplasts. More specifically,
chlorophyll resides in the thylakoid membranes. The chlorophyll absorbs energy from
sunlight, and it is this energy that drives the synthesis of food molecules in the chloroplast.
The chloroplasts make food via photosynthes is, and veins in the leaves and other
photosynthe tic parts export sugar to the roots and other non-photosy nthetic parts (parts that
have no chloroplasts and thus are not green) of the plant. Photosynthet ic pigments act as light
receptors. The process of photosynthes is depends upon the efficient capture of light energy by
aggregates of pigments present in photosynthetic tissues. It is know that as light comes into
contact with matter, it may be reflected, transmitted, or absorbed. ~~~ ances that selective!_y
~~sorb visible light are called pigments. Different pigments absorb light of different
wavelengths , and the wavelengths that are absorbed disappear. This is important because light
has to be absorbed before it can be of any use in a photobiological reaction. The pigments of
chloroplasts absorb blue and red light most effectively, ~nd t_ransmi_t _or_~eflect green ligb~-
~ hich is why leaves appear green.
~ Chlorophyll is a complex molecule. All chlorophylls have a lipid-soluble
~ ~ophobic) hy_Qrocarbon tail (C20H39 -), a flat hydrophilic head with <!__magn~~~~~~n at its_

-
centre; while different chlorophyl1s have different side-groups on the head. The tail and head
.
are Jinked cogether by an ester bond. Wh_ile the hydrophobic tail projects
the hydrcphilic head acts as solar panel on the thylakoids.
into the thylakoids,

~, he arrangement of the chlorophyll and other pigments in the thylakoid~ is termed

Photos/ stem. Several modifications of chlorophyll occur among plants and other
photosynthet ic organisms. All photosynthetic . organisms (plants, certain protistans,
prochlorobacteria, and cyanobacteria) have chlorophyll a. Accessory pigments absorb energy
that chlorophyll a does not absorb. Accessory pigments include chlorophyll b (also c, d, and e
· in algae and proJ.istans), xanthophylls, and carotenoids (such as beta-carotene). Chlorophyll a
• absorbs its energy from the violet-blue and reddish orange-red wavelengths, and little frorTl
the intermediate (green-yellow-orange) wavelengths.
Carotenoids and chlorophyll b absorb some of the energy in the green waveleng th ·
Both chloroph yll s also absorb in the orange-red end of the spectrum (with longer waveleng th5
and lower energy ). The origins of photosynthetic organisms in the sea may account for thiS-
Shorter wa velengths (with more energy) do not penetrate much below ..5 meters deep in sea
water. The ability to absorb some energy from the longer (hence more penetrating)
 wavelengths might have been an advantage to early photosynthetic algae that were not able to
be in the upper (photic) zone of the sea all the time. The action spectrum of photosynthesis is
the relative effectiveness of different wavelengths of light at generating electrons. If a
pigment absorbs light energy. one of three things will occur. ~nergy is di ssipated ash~. ~
~ergy nlll}'... be e!!litted im~ <li ute!}, us a J onger ~ avelength , a uhcnom£!!2!1 know!!..-A
~ ; orescen£e. Energy muy trigger u chemical reaction, as in photosynthesis. Chlorophyll only
triggers a chemical reaction when it is ussociated with proteins embedded in a membrane (as
in a chloroplast) or the membrane infoldings found in photosynthetic prokaryotes such as
cyanobacte,i a and prochlorobactcria.

0= --CH
I

\
('
( '-t .

(
I •
H
0
//
C
,, '
()
0
Chlorophyll a: 3
,. . . . H
I
0
I
•,: ll ' 0. ,
Chlorophyll b: . ~ = -C
~o
(H

( 'i ( 1-1
\ /
C
I
r 1

Fig. 2.7: The molecular structure of ch,lorophyll (Mg is at the contre of the head)

Photosynthesis in bacteria and algae.

Photosynthesis does not only occur in the higher green plants~ it occurs in algae and
bacteria as well, albeit a little differently. Bacteria are unique in that they do not have
chloroplasts, which, as we know by now, are the main sites of photosynt'-esis in the green
Plants. They utilize different pigments, and sometimes their products differ significantly. For
~x.amplc, modern archaebacteria photosynthesfze witi!_ t~e_eigment_bactcri..orbodopsin (which,
Ythe way, is structurally related to the visual pigments in our eyes). Green and purple sulfur
!actena o~c their respective colors to_!l~etiochlorQithY!l, their rAain photosynthetic pigment.
lants which perform green plant photosynthesis as opposed to bacteriar photosynthesis have
c_hlorophylls as common pigments having major absorption peaks in the visible region of the
light spectrum. Bacteriochlorophylls, however, absorb near · infrared radiation; thus,
Photosynthetic bacteria grow in regions where higher plants cannot.
 "° • ()
.
I
)
". }_ ll \
' IC' 1) ' { (·i

Chor ophy lls °' r Z - r ,/

· c hi orop hYII ·a All photo synth_ etic
_ or e-0 an1 srn s
·
One very important chloro phyll ,s
photo syn th ell~ pigment. It is
(exce pt a few group s of bacteria) have this as their main
reaction_s. which conve rt solar
impo rtant becau se only it can participate direct ly in the light
the only pigm ent in chloroplasts
energ y to chemiciil energy. But chlorophyll a is not
th
light and transf er e energy to .
imixm ;nt -lo photo synth ;~s. Other pigme nts can absor b
refer ~ ~~~ "" J~er pigments as
0
chlor ophyl l a, which then initiates the light reactions. We
ophyl l, chlorophyll b. It is very
acces sory pigme nts. One of these is anoth er form of chlor
has its own uniqu e color. The
simil ar to chlorophyll a, but different enoug h so that each
point is, if chlor ophyl l b absorbs
forme r is blue- green , while the latter is yellow-green. So the
the energ y lo chlor ophyl l a, which
a photo n of sunlig ht (discrete packet of energ y), it conve ys
other chlor ophyl ls exist along
then acts as if it had directly absorbed the energ y itself. Many
ophyl l c is prese nt in the brown
with the two we have just emphasized. For exam ple, chlor
nt along with chlorophyll a in
algae , the dinof lag~l ates, and the diatoms. It is gener ally prese
is chlor ophyl l d, which is almost
most marin e photosynthetic organisms. Anoth er exam ple
ophyl l a.
exclu sive to the red algae; it acts as a minor pigme nt to chlor

Carotenoids
other
Well, what 'take
. ...,.
s leaves orange and yellow ? The chlor oplas t also has many
There are a group of pigments
accessory pigme nts that aid in the process of photosynthesis.
shade s of yellow and orange.
called carotenoids, which unlike chlorophyll , are different
thylakoid memb rane. Carotenoids
Carotenoids reside with the two types of chlor ophyl ls in the
of light that chlor ophyl ls cannot.
are important because they can absorb certain wavelengths
and yellow. Carot enoid s are also
These are the structures that cause leaves to appea r orang e
photo prote ction. Excessive light
iI!!PQrtant ~ca~ they ~e involved in a function known as
d of transm itting energy to
intensity can damage the c hlorophyll pigme nts~ so instea
energ y from chlor ophyl l, thereby
chlorophyll, some carotenoids use photoprolection to accept
found in the photosynthetic
protecting them from harm . A variety of carote noids are also
bacteria.

Xanthophylls
Carotenoids are divided into· two classes: carotenes and
xanthophylls. The major
and brown algae . The main
x.anthophylls of higher plants are also present in the green algae
al, is called fucoxanthin, which
x.anthophyll of diatoms, in specific, and brown algae, in gener
.anthin, howe ver, is not present
is what gives these organisms their brownish color. Fucox
the dinof lagell ates (diatoms).
out!ide the kingdom Chromista (brown algae and relations) and
divisions. The important point is
Several other x.anthophylls are also prevalent in other l}lg~I
unique colors.
that the different x.anthophylls are what gi ve some algae t!;i.~i~ ,
Phycobilins
and carotenoids. are 1he
A final group of pigments, in addition to the chloroph yll s
arc rather widely distributed
phycobilins . Unlike the chlorophylls and car9tenoids, \'. hich
ution and are found only ill
amon g various plants, phycobilins have a relativel y narrow distrib
pigment helonging to this fornil Y.
the red algae and cyanobacteria. Phycoerythrin, an accessory
is what makes red algae commonly red. Conversely, phycocyanin, another accessory pjgment,
is what causes the cyanobacteria to appear blue-green. The phycobilins and other accessory
pigments are what also make possible the absorption of filtered blue and green wavelengths in
deep waters.

"'
.&:.
-~;
-
I
I
a,
C
800 _,0
J
106

Fig. 2~8: The electromagnetic spectrum

The chloroplast
The chloroplast is the organelle of photosynthesis in eukaryotes, .its number may vary
.Rb_yll cell of a higher plant. It
from one in Chlorella to more than a hundred- iQ.lliili.£ade...m.eso
----- _,__ _
is usually 3-l0µm in diameter, and visible under light microscope. In many ways, the
chloroplast resembles the mitochond.rion.
Both are surrounded by a double membrane with an intermembrane space.
Both have their own DNA .
Both are involved in energy metabolism. ' .
Both ha~ membrane reticulations fillin~ their inner space to increase the surface~
area on which reactions with membrane-boun d proteins can take place. The chloroplast ~
three membranes: inner, outer, and thylakoid. It has three compartments: stroma, thylakoid
1>8£e, an_d}nter-mem~r!ne sp~~
_8 These compartments and the membranes that separate them
serve to isolate different aspects of photosynthesis. Dark rcactibn...§.. take place in the stroma,
~ _Q!l the thyl~~~id IJ!embranes. The thylakoid is the s~tural
-Ught reacli<?JlS take pJace
unit of photosynthesis. Both photosynthetic prokaryotes and eukaryotes have these flattened
&acs/vesicles containing photosynthetic che,micals. Only eukaryotes have chloroplasts with a
sulTounding membrane. Thylakoids are stacked like pan~akes in stac_!!!oa~n collectjye~ <

~ The areas betw~~ g;:r;a are referred to as stroma. While the mitochondrion has two J

fllembra~e systems, the chloropl;t has three, forming three compartments. • ·

~ ,\\ (i\~,:,tl - '/ (' \ ,o ,{\_ \ \11 '-;J ~ Y," •- < l \ l <" ~~
 >) -.:
'

Granum
(stack of
thylakoids)
lntermembrane ----+11 1•1111 \
spa

Inner l r' I
I

mem~rane - \ " ~ _T _
. .I
Outer ~ " ,l l ~ /
membrane , :. ,, •• . .
"· .. -<'· !1" . "

Fig. 2.9: The structure of a chlorop last.

Stages of photosynthesis
Photosynthesis is a two stage process. The first process is the light depend ent process
es
(light reactions), which requires the direct energy of light to make energy carries molecul

ction
--
tha re used in the second process. The light independent_process (or dark
are used to form C-C covalen t
---- reactions) occur~
bonds of_carboh
___,
ydrate~:
~hen the product s of t~e light..r.ea
process
The dark reaction s can usually occur in the dark, if the energy carriers from the light
y
are present. Recent evidenc e suggests that a major enzyme of the dark reaction is indirectl
ligh!
stimula ted by light, thus the term dark reaction is somewh at of a misnom er. The
asts.
t. - and-the-dark
reactions occur in the -grana - --of--the chloropl
- reactions take place- in-the stroma ______,..--,
~~ l~-:. \)~rt, . .·
·. h .· "'11 ¥ ~ r •. j
L 1g t reactao ns • · ,._:J · • .·

In the light dependent processes (light rea~tions), light strikes shlorop hyll a in such a
is
way_as to excite electrons to a higher energy state. In a series of reaction s the energy
in the
convert ed (along an electron transport process) into ATP and NADPH . Water is split
used to
process, releasin g oxyg~. as _a b-y-prbduct of the reacti~A. l'he AtP and NADPH are
the sun
make C-C bonds in the light independent process (dark reactions). Light energy from
After
is co·~~~rted into u~eful. chelT\ical to~ls through the light reaction s of photosynthesis.
of its
light is absorbe d in the chlorop lasts by the pigments, the pigmen t will boost one
and then ~
electron s to a higher energy level. .Of course, a lone pigmen t would simply excite
bYc
give off its energy as heat resetting itself for another absorpt ion. This process is replaced
sts,
a 1
gatherin g several pigments togethe r to form an attenna e comple x within the chloropl
it at tht ,
which is similar to a· funnel that gathers water from a large area and concentrates
a. Th' ,
center. At the center of this comple x will be one pigmen t, most likely, chlorophyll
fro~"
position of this pigmen t allows it to receive all the ene.rgy the other pigmen ts received 1
th at ' '
the light and thus excite one electron to a very high energy state. This is the energy
cycle,~:,
used to power the breakdo wn of carbon dioxide into nutrition for the plant in Calvin 1
up ·
dark reaction s. The combin ation of the antenae comple x and the reaction center make <I
•
photosy stem, the light harvest ing comple xes of the thylako id membra ne.
tt 0 + light energy -,->½ 02 + 2H+ + 2e- (Photolysis or Hill's reaction)
2
 , hl' rn,J uc l nr phnt, 1synthesi s. L·ould on~1n
,1\,. - ~l' ll
~
atc fr 0 m c iih c r !he CC) or. t 1l C 'f
: r ~ 0 Siurt ing
uinal L'o miio unds l d h
:l'omrciunJ
..
<., _ To determin e " h1c h of th ese · ori ~ ~ con I n 1u1 c to t c 0 2 end
· 1>roJuct. an isotopi c tracer e'Zpcrim e nt was performe d usin g '' O:

180 . h .
0 1s a ca ,·y isoto pe of oxygen
0 H.:t o + CO 2 ~ ie lds I R0 2
1
o H20+C R0 2 yie lds 0 2

Therefore . the 0 2 end product must originate from water and not fro m the carbo n dioxide .

~~\<- ~ X.~'
In the light independ ent process, carbon dioxide from the atmosph ere (or water for
aquati c/ma,ine organism s) is captured and modified by the addition of hydrogen to form
carbohyd rates (general formula of carbohyd rates is [CH 2O]n)- The inco1por ation of carbon
dioxide into O.!:._g anic compoun ds is known as carbon fixation. ,The energy for this comes fr~~
,.....__ -- --
the first phase (light reaction) of the photosyn thetic process in the fo1111 of ATP and NADPH.
Li ving systems cannot directly utilize light energy, but can , through a complica ted series of
reacti ons, convert it into C-C bond energy that can be released by glycolys is and other
metabolic processe s. 9. l 0 2.. -llt-t:>.- ->2C f+;i._ ~ -+ ~

Photosystem
Photosx_s tems are _aJTan~~ ent~ of chlo roph yll ~ other pigments _packed into
~koid s. Many Prokaryo tes ha ve only one photosys tem, Photosys tem 2 (so numbere d
because, while it was most likely the first to evolve, it was the second one disco vered).
Eukaryotes have Photosys tcm 2 plus Photosys tem 1. Photosys tem I uses chloroph yll a, in the
form refen-ed to as P700. Photosys tem 2 uses a fo1111 of chloroph yll a known as P680. Both
"acti ve" fo,ms · of chloroph yll a function in photosyn thesis due to lheir associati on with
proteins in the thylakoid membran e. The photosys tem has antenna complex , containin g 200-
300 pigments , each collectin g light of differenr waveleng ths thereby making the process
efficient. -p s- 1 - p~
t>S t\ := ~ b~
Photophosphorylation
Photopho s phorylat!Q!l is the pr~ ss of convertin g_elJ£!_·gy from a li g_ht_:~xcit ed electron
t

_!!l to the pyrop~s phate bond of an ADP molecule . This occurs when the electrons from water
• -

are excited by the light in the presence of P680. The energy transfer is similar to the
~ cherniosmotic electron transport occurrin_g in the mitochondria. Light energy causes the
" removal of an electron from a molecule of P680 that is pa,1 of Photosys tem 2. The P680
e
req uires an electron , which is taken fro m a water molecule , breaking the w ater into H+ ions
,e . .anu o-~ions. These 0 ·2 ions combine to the diatomi c O ] thal is released . The electron is
fo1111

,;i . • "boosted" to a hi gher energy state and attached lo a primary electron acceptor.
which begins a
11
1· 5
. ·Jle ri i.:~ of redo x reaction s, passing the el ectron through a se ri es of e lectron ca1Tie rs. eventua Y
d , tlltac hing it lo a mo lecule in Photosys te m l. Light act s on a molec ule o f P700 in PhotosyS le rn
~[{g)_ ca11-; ino an e lec tron tn he "boosted" to a still hi ghe r potential. The electro n is au ac hcd 10 a
mtdij le, i.: 111 ; r, mary l'lcLI rnnacce ptor (th at is a differe nt mo lec uk from the nnc assoc iated wi th
 REDOX
POTENTIAL
(V) \
1

photon e- j
~ 1
L_ I
' / e-
@ -•~-✓/

DIRECTION OF ELECTRON FLOW

Fig. 2.13: Cyclic photophosphorylation

Dark reactions (Cal"in cycle) #

Melvin Calvin was a Professor of Chemistry at the University of California a!
Berk.eley when he and his colleagues eluciuatcd the pathway of carbon fixation into glucose.
Cal vin "'as able to determine the path of carbon by exposing at various times and
1
illuminations, the ~nicellular alga Ch/ore/la to °'C02. a radioactive isotope of carbon. Then.
they quickly dropped the alga into a pot of boiling alcohol to stop the process but still
preserve the labeling of the isotope. Calvin and his colleagues were able to detenninc 1hal
fixation of carbon occurred in three stages that make up one cycle. Calvin was awarded 1he
Nobel Prize for Chemistry in 1961 for this work The staoes •tre pres t db ·
.
0
' en e e 1ow:
Stage J: A very slow enzyme named Rub1sco takes ·m inc • CO I -nd 1
ommg 2 mo Iecu e , '
. < 1
attaches it to a five carbon sugar calleu R1bulose Bi sphosplnte (R BP) R . . h fi rst \
. . • c u . ub1sco 1s t e 1
step and is just an abbrev1at1on for nbulose bisp/wsphare ca b . l I ihi5 \
r OJ:y ase. .Unfottunate
. .
Y ~ •~
enzyme is very slow and thus the plant counteracts this hy hav·
0
.
abundant enzyme on earth. The 6-carbon sugar formed from th · . of. 1t ' as 1t ts the ni. nJ ,
mg a lot
. . . . called .3-Phosphogly
smalle,. units is union ts very unstable ·1-f
breaks into two 1dcnt1cal
. .
of photosynthesis cerate (3-PGA). the first prodU'

Stage 2: ·11 , , , ~,age mostly a series of h .

is 1,
. c em1cal . . .
ieactJons t at th' he .
Phosphoglyceral e ~ocs throu gh. where ll picks up anoth .
1
c, P iosrhat e ion. , anc..l two c Iec rro~:-
from NADPH. hnally. it ends with a new name and ne , . ·
. I1Yde .
Phosphate (G3P) . Remember. ,ve ha ve two of these, onewt sti ucture c·..t II c·<.I GI yceralde
ravels off to form glucose wh1·1e t 1
other re-enters the cycle in the sta~e 3. So. it takes two t
. ~ urn s of the , I . ,,
molecule of glucose. c ye em order to create l
Cellular Respiration
Cellul ar ~spirntion is t.he procc1,s 9 f oxidi ~i!rn food molec ules, like glucose to carbon
of ATP
dioxid e and water to gene rat_c energy . The ene rgy re leased is trappe d in the form
(adcno sine triphos phate ) for use by nil the energy-consu ming activit ies of the cell.
ATP
It 1s culled adenos ine tripho sphate. J!
is the energy curren cy of...the cell , and it is
is added to
formed hy th pn'X·css of phosph orylati on, whereb y an inorga nic phosp hate gro up
of water
ADP (adcno sme dtphos phate) . Hydro lysis of a molec ule of ATP in the presen ce
yields 30.6KJ
ADP + Pi + energy -+ATP + H20
ATP+ H 20 -+AD P + Pi + 30.6 KJ
It ,s the cells· universal energy carrier , just like a battery , supply ing energy for wide
of foods
ranges o f act1 vuies. ATP can be rechar ged using the energy from oxidat ion
facilita ted
(oxida n"e phosph orylation) or light energy (photo phosph orylati on). Its synthe sis is
c base
by enzym e A TPase . Structu rally, jJ is IJlade of a ribose su~ (5-car bon), an organi
(adcomc) and three phosph ate groups . ,,
NHt
N I

_
0
II
0
II
O-P- 0-P- O-P- 0-CH t
0
II
<D N

adenine
J- J- J-
H
OH OH
ribose
Fig. l. J8: The structu re of ATP

Cellul ar respiration invol ves oxidation of foods such as carboh d . t· d 1·1s

While carhoh ydratc~ arc fi n,t hydrol yzed to monos.acchan_des and Y rates,
· · I
ats an
· the
° •

proces s. fat s c:1nd Q!<Hcin s ~an be s d durin the . pnnci pa ly used tn

nod of starva f Th irt
.;::;;. •on. ~,r,roc ess occurs
two phases ; glyco lysib and Krebs c yc le.
,,
- \l, '
 ( ; l~:<'ol ysis
/ T here are two important ways a cell can harvest energy from food: fermentation and
'Sdlular respiration . Both start with the same first step: the process of glycolysjs ¼hi.ch ,s the
hreakgo wn Qr splitting of glucose (6 carbons) into two 3-carbon molecules called pyruvic - -
, -----=::::::========-=-=-=~::.!_~ =-...:~=-=~=-=-~_:_:_:_:_:._ ~--
aci<J... The energy from other sugars, such as fructose, is also harvested using this proces~..
G %
Glycolysis is probably the oldest known way of producing ATP. There is evidence that the
process of glycolysis predates the existence of 0 2 in the Earth ' s atmosphere and organelles in
cells. Glycolysis does not need oxygen as part of any of its chemical reactions. It serves as a
first step in a variety of both aerobic and anaerobic energy-harvesting reactions. Giycol&
occurs in the cytoplasm of cells.; not in some specialized organelle. Gl:xfol ysis is the onl y
•
metabolic pat~ay founc! i!!_ all living orga~ism~. £
Glycol ysis is a three step process: the first is the priming phase requiring energy in the
form of ATP phosphorylation of sugars. The second is the lysis or splliting of the
phosphorylated 6C sugar to 2 molecules of 3C sugars, while the third phase is the energy
'
yielding phase thn,ugh the process dehydrogenation.

Glucose
ATP
hexokin11• ,
gtucoklnllt
ADP

Glucos7~hosphate
ph01phohe10s1
l1onera■e,

Fructos -phosphate
ATP
pho,pho-
fructokinase-1
ADP

Fructose-1,6-blspho&phate

Glyceraldehydjo-3-phosphate •t-rio-■-tpho-,-p-ha-t•-Dihy~=one
tsomerase

Fig. 1.19: Glycolytic pathway showing the first and second phases of reac rion
 Glyce ralde hy;de -3-ph ospha te
NAO'+ P,~gl ycera ldehy de--3 .phos phate
dehyd rogena se
NADH+ ..r

1,3-b ispho s hogly cerat e

ADP
phospho glycer at•
kin as•
@
3-phosp1oglycerate
phosp hoglyc erate
mutase
2-phosJll'!oglycernte
l ~nolace

~3r;,;~e
Pyruvate

te
Fig. 1.20: The third phase of glycolysis showing the formation of pyruva

sphate . In
In the first phase, 2ATPs are used to convert glucose to fructose-1,6-dipho
molecules of of 3C
the second phase, fructose-I, 2-diphosphate is degraded to produce two
hates are degraded to
phosphates which are isomers. In the third phase, the 3C sugar ptiosp
The energy balance is
pyruvate, with the production of 4 ATPs and 2 equivalents of NADH.
and 2NADHs are
such that two molecules of ATP are used, and four molecules of ATP
.les of pyruvic acid.
produced by I molecule of glucose that is oxidized to two molecu
e. Pyru vate is the ,
Therefore the net energy of glycolysis is 2ATPs per molecule of glucos
state of the cell i.e. the
branch point molecule in glycolysis. Its fate depends on the oxidation
availability of oxygen or otherwise. When oxygen is not available or when g.tycolysis is
~ J lolln) ,1 ,, \I
and skeletal muscles
proceeding at a high rate, or in anaerobic organisms, or in erythrocytes
lactic acid through the
under conditions of exe,tion, pyruvic acid is converted to ethanol and
pyruvate enters the
process of fermentation. However, when oxygen is available,
which is the product
tricarboxylic acid (TCA) cycle or Krebs cycle in the form of acetyl-CoA,
etely oxidized to CO2
of pyruvate dehydrogenation reaction. In this way, pyruvic acid is compl
n called aerobic
and H20, liberating large amount of energy , in a sequence of reactio
respiration.

produc t, and a
Fermentation , pyruvic acid molecules are turned into some "waste"
e actually four are
little bit of energy (only two ATP molecules per molecule of glucos
many possible types of
produced in glycolysis, but two are used up) is produced. Out of
acid fermentation and
fermentation processes, two of the most common types are lactic
\
alcohol fermentation .
 ·r :r l"\t•~' .
I xt,l· .ll' HI kr111L·111a111111 1s earned lllll h) sn111c fuJJg•. so~ ~~ lrkc thL·

/ ,1rtol1un'l/11s acitloplti/11.,· 111 y1)g11r1 , / i11 ·robol·il/us sn in fc nnc_nt~.d.J~ J~ sud1 a~ (1 g1. g,1rn .
1111 . 111111 t'l l' . :rnd s1Jllll't1 111cs b} our musl'lcs. N,>rrnull)' our muscles dn cellular rcsprratron li ke
1

tilt' 1\·st ,,t ,1111 b\ 1d1t·s. 11s111g 0 1 s11pplinl by t>ur lungs and blood . I lowcver. under greater
nett HIil "lie n the· 11,ygc11 suppl~cd by the lungs and hloou system G111 ·1 gel there fast enough
11, i...t·c·p 11p with the 11u1srks · lll'L'ds. t)lff' musc les r an switch mer anJ do lactic acid
fl'rt m·111 :,t1 <111 . l11 the pmn.· ss lll' laL'lir acid fermentation. the J -carbon pyruvic ac id molecules
:11\ ' 111111cd 111t "~1c.:!.kJ).cid . It is the prcscnt·c nf lac ti c acid in y9g!!f1 th ~t ,gives it its sour tas .
uml 11 1s the p,~sencc of lactiL' ncid in ('Ur nmscks "the morning after'' .
that makes them so
'-

:-lit\: . Onrc our rnuscks form lactic aciJ. the y cu11·1 do anything else with ,t. so until it is

e.rmJunll y ,, aslwd awny by the hloL1d sir-cum and earned lo the li ver (which is. able lo get rid of
~

11). \Htr t) \'l'r CXl'l1Cd mu:-:c k s feel stiff and sore even if they ha ven' t been ph ysically injured.
G h,cost· - pyruvic acid -+ laclctic acid+ 2 ATPs
I
'c.. 11 • C.
Akohol fc1111entatiun is cnnied out bv yeast and some kinds of bacteria such as • - - - -- - - -
..... - • 4. •

/ \'111m11mws mobili.,·. The ·•w.,stc" products of this pnx-ess are ethanol and carbon di ox ide
(l'l)~). I lumans have long taken advantage of this process in making bread. beer. and wine. In
hrcud making. it is tht~ CO~ which fonns and is trapped between the gluten (a long protein in
wlit·:11) llllllt"t'tilcs that causes the bread to ri se. and the ethanol (often abbreviated as EtOH)
n:ipt,rntmg that gives it its wonderful smell while baking. The effects of 1hc e thano l in beer
and something with which many college students arc familiar and it ,s the _£Q;.
\\' tnC arc
erndurcd by the process of fermentation that makes these bevera~s effcrvcsccnl~

Glucose-+ pyruvic acid -+ ethanol +CO~+ 2 ATP ~

I \ '-I'
Krebs cycle
') ltj',' ~ ~v( .c..rrc\j (. t \,

The pyru vate molecules ·produced during glycolysis contain a lot of energy i11 the
bonds between l.hcir molecules. In order to use that energy. the cell must conven it into the
fom1 of ATP. To do so. pymvate molecules are processed through the Krebs Cycle. also
~ known ~ t.he eitrir acid cycle. The cycle was elucidated by Sir Hans Krch;-hence its name as
th
~ Kn!bs cycle. Since rhe acids that a[e_futl]led~durin ~ the prucess _c_xist as salts, their names
al~vuys end ,virh-ate e.g. ,..si t.p ile (11m;1rate e~c ~ The proce~s occurs in th; ..~atrix of the
mHochondrio~been catalyzed by a nulll.ber o[ en_zyrne~. Because glycolysis produces two
P~vate molecules from one glucose, each glucose is processed through the Krebs cycle
twice F . . . . +
· or each molecule ot glucose, six NADH2 • two FADH1 • and two ATPs are produced.

lnn...-
•mbrane

Malrix
I •
C

I ntermembra ne
space
 t·· JAOH++ H+·~- ----,
II.AD+ _ _ _ __,,
8

~
7 3
F A.DH,4
. ;___ _

FAD--_ _,,

GTF'

Fig. 1.22: Ttw TCA or Krebs cycle

1. Ptior to entering the Krebs Cycle, pyruvate must be converte d into acetyl CoA
pronounced: acetyl coenzyme A). This is achieved by removin g a CO 2 molecule
from pyruvate and then removing an electron to reduce an NAO+ into NADH. An
enzyme called coenzym e A is combine d with the remainin g acetyl to make
acetyl CoA which is then fed into the Krebs Cycle. The steps in the Krebs Cycle are
summarized below:
2. Citrate is formed when the acetyl gr~up from acetyl CoA combine s wi th
oxal~acetate from the previous Krebs cycle.
3. Citrate is converted into its isomer isocitrate.
4. lsocitrate is oxidized to form the 5-carbon a.-ketoglutarate. This step releases one
molecule of CO2 and reduces NAO+ to NADH/ .
5. The a -ketoglutarate is oxidized to succinyl CoA, yielding CO2 and NADHi°'·
6. Succinyl CoA releases coenzym e A and phosphQt'Ylates ADP into ATP.
7. Succinate is oxidized to fumarate , converti ng FAD to FADH2.
8. Fumarate is hydrolyzed lo form malate.·
9. Malate is oxidized to oxaloacetate, reducino0 NAO+ to NADH /.
~
Elertron transport chain
What happens to the NADH d FAD + ·
an 2 H 2 produced dunng the Krebs cycle? The
molecules have bee~ reduced, receiving high energy electrons from the pyruvic acid
molecules that were dismantled in the Krebs Cycle. Therefore, they represe-nt energy available
to do work. These carrier molecules transport the high energy electrons and their
accompanying hydrogen protons from the Krebs Cycle to the electron transport chain in the
inner mitocho nd rial membrane. In a number of steps utilizing enzymes on the membrane,
NADH/ is oxidized to NAD+, and FADH2 to FAD. The high energy electrons are transferred
to ubiquinone (Q) and cytochrome c molecules, the electron carriers within the membrane.
The electrons are then passed from molecule to molecule in the inner membrane of the
mitochondrion , losing some of their energy at each step. The final ~ransfer involves the
combining of electrons and H2 atoms with oxygen to form water. The molecules that take part
in the transport of these electrons are referred to as the electron transport cha.in.
The process can be summarized as follows: the electrons that are delivered to the
electron transport system provide energy to "pump" hydrogen protons across the inner
mitochond1ial membrane to the outer compartment. This high concentration of hydrogen
protons produces a free energy potential that can do work. That is, the hydrogen protons tond
to move down the concentration gradient from the outer compartment to the inner
compartment. However, the only path that the protons have is through enzyme complexes
within the inner membrane. The protons therefore pass through the channel lined with
enzymes. The free energy of the hydrogen protons is used to form ATP by phosphorylation,
bonding phosphate to ADP in an enzymatically-mediated reaction . Since an electrochemical
osmotic gradient supplies the energy, the entire process is referred to as cherniosmotic
phosphorylati on. Once the e lectrons (originally from the Krebs Cycle) have yielded their
energy, they combine with !xygen to form water. If the oxygen supply is cut off, the electrons
and hydrogen protons cease to flow through the electron transport system: If this happens, the
proton concentrat.ion gradient will not be sufficient to power the synthesis of ATP. This is
why we human beings, and other species, are not able to survive for long without oxygen!

lntennembrane Space
:c
:c :c l"'t ..., •
I + °'a I
,l ♦1 +i . :
('I 0

Matrix
 based on
. hi ornetrY,
sto1
The amount of ATPs produced in cellular respiration is not c generated by
h electron
three reaons: each electron generated by NADH generates 3 ATPs, eac . . cid yields 2
. . . .d tO succin1c a
FADH2 yields 2 ATPs, while the conversion of a-ketoglutanc act y yield in
th
ATPs. Glycolysis, yields 2 ATPs per molecule of glucose oxidized. Thus e energ
a complete oxidation of a molecule of glucose can be summarized thus:

10 (NADH) yield 30ATPs

2 (FADH 2) yield 4 ATPs
Conversion of ketoglutaric acid to succinic acid yields 2 ATPs
Glycolysis yields 2 ~ T s
TOTAL: 38 ATPs
,,
•
Oxygen debt and exercise
ATP is not a storage form of energy but a 'curre~ futf that can liberate energy
when needed almost instantenously by the living cell. Other macromolecules such as sugars,
proteins and fats are forms through which energy can be stored. So, ATP is only available in
little quantity in the body. Sudden change from rest to exercise requires a lot of
. 0 2, to
generate energy by aerobic means. In. this .
regard, 02, may not be sufficient to~
. ....arry out the
P rocess. Therefore, lactate fermentatlOI) 1s s~1tched -~n to _gel_!erak__AIP_at a nfu., .....
- ----- - - --~ ~ _ [ _rate (2 5
times faster). Also, glycogen can serve as source of glucose for the process. - ·