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Glossary

A Hyperfine interactions between the electron spin and nuclear spin. acetylcholine receptor channel A type of receptor for the
A DNA A conformation adopted by dehydrated right-handed neurotransmitter acetylcholine, found on skeletal muscle cells
double-helical DNA. The molecule is shorter and wider than a B and some nerve cells. Also known as the nicotinic acetylcholine
DNA of an equivalent number of base pairs, and the base pairs receptor, it is made up of a number of protein subunits which form
are tilted relative to a line perpendicular to the helical axis. an ion channel through the cell membrane. The channel is opened
A site The acceptor (A) site on a ribosome at which the incoming by the binding of acetylcholine to extracellular sites on the protein.
aminoacyl-tRNA binds and is decoded. The site involves both the Achaete-scute complex (AS-C) A set of four closely linked genes
small and large subunits and binds a complex of elongation factor in Drosophila that are important for the formation of the nervous
(EF-Tu in bacteria; EF-1 in archaea and eukaryotes), GTP and system. They encode transcription factors of the basic helix-loop-
the aminoacyl-tRNA. helix (bHLH) type.
abaxial Facing away from the main growth axis. For example, the achene A single-seeded dry fruit that does not split open.
underside of a leaf is the abaxial surface, as it faces away from the Acheulian Stone tool industry dominated by handaxes.
direction of growth of the stem. aciclovir Nucleoside analogue inhibitor of human herpesvirus 1.
ABC proteins Proteins containing the ATP-binding cassette motif. acid A substance that donates protons (H 1 ions) to water (forming
They include several types of transporters. H3O 1 ) or to another acceptor. An acid can also be considered as
abiotic Of or pertaining to the nonliving. a compound that dissociates to produce H 1 and its conjugate
abiotic factors The chemical and physical environmental factors in base.
ecosystems. acid–base catalysis Enzymatic catalysis that involves a general
acid (a molecular species that can donate a proton, e.g. histidine)
absorbance optics Optical method for measuring the concentra-
and a general base (a molecular species that can accept a proton,
tion of a substance by measuring the loss of light due to
e.g. aspartic acid) at the enzyme active site.
absorbance of photons by molecules.
acid hydrolases Hydrolytic enzymes that have maximum activity
absorptive endocytosis Internalization of a protein by a cell after
at acid pH. In animal cells they are found mainly in lysosomes.
the protein has bound weakly and nonspecifically to charged
acquired immunity See adaptive immunity.
groups at the cell surface.
acquisition Uptake of virus by a vector.
acaricide A chemical treatment applied to animals to kill ticks and
acquisition period Time required for a vector feeding on the host
prevent transmission of tick-borne pathogens.
to acquire the virus.
accelerated run The propulsive run a large bird or a poor flier acrocentric A chromosome with the centromere near one end and
needs to make during take-off in order to achieve the minimum thus having one very short arm and one long arm.
flight speed.
acron The most anterior portion of the crustacean body, generally
acceptor The larger of two reactants that interact reversibly to thought not to be homologous to the other segments and usually
form a complex. including the protocerebrum and eyes.
acceptor arm The end of a tRNA molecule to which an amino acid acropetal Refers to leaves, flowers, or other structures developing
becomes bound. It contains both the 50 and 30 ends of the tRNA. successively along an axis so that the youngest are at the tip.
The 30 -terminal sequence of cytidine-cytidine-adenosine (CCA) acrosome A cap-like structure covering the anterior part of the
overhangs the end, and the terminal A is the site of ‘acceptance’ of sperm head, containing enzymes important for fertilization.
the amino acid. actin The protein monomer (G-actin) that polymerizes to form the
acceptor end Four nucleotides at the 30 end of tRNA, which microfilaments (F-actin) of the eukaryotic cell cytoskeleton and
form the site at which the amino acid becomes bound. The last the thin filaments of muscle and other contractile structures in
three nucleotides are cytidine-cytidine-adenosine (CCA) and the eukaryotic cells.
ribose of the terminal adenosine forms a bond with the amino actin-binding proteins Proteins that interact with actin, either to
acid. modulate its polymerization and depolymerization, or to link
acceptor stem That part of the acceptor arm of a tRNA molecule actin microfilaments to each other, to membranes or to other
in which nucleotide sequences from the 50 and 30 ends are paired to proteins.
form duplex RNA. actin cycle The polymerization/depolymerization of actin fila-
accommodation An increase in the threshold for an action ments, which is coupled to nucleotide hydrolysis and exchange.
potential that occurs in some neurons during a slowly developing actinosporean phase The alternate phase of development in
or prolonged depolarization. The result is that only a few action myxozoan parasites, which occurs in annelid worms and
potentials are generated during prolonged depolarization above culminates in the formation of actinosporean spores.
the normal threshold level. actinotroch, actinotrocha The larval form in the Phoronida, a
Ac/Ds Activator and Dissociation, the two mutable loci originally phylum of marine worm-like animals.
identified by Barbara McClintock that were able to move in the action potential An electrical impulse that propagates along
genome. neurons, and other electrically excitable cells such as muscle, in
acetylcholine A chemical neurotransmitter that is released from response to stimulation above a certain threshold. It consists of a
the ends of certain types of nerve fibre when they are stimulated. It self-propagating depolarization of 0.1 V in the cell membrane and
transmits a signal to an adjacent nerve or muscle cell by binding to carries an informational signal from the cell body to the nerve
receptors on the target cell surface. terminals, where it activates synaptic transmission.

143
Glossary

activating regions Sites on transcription-activating proteins that additive distance The distance between two operational taxo-
make direct contact with RNA polymerase during the initiation nomic units in a phylogenetic tree, which should be equal to the
of transcription at gene promoters. sum of the connecting branch lengths. This assumes a constant
activation Stimulation of a cell, protein or gene to carry out its rate of substitutions since the time of divergence from the
function. A gene, for example, is said to be activated when it is ancestral sequence.
being transcribed; an ion channel is said to be activated when it is addressins Proteins on the surface of endothelial cells which act as
open. ligands for lymphocyte homing receptors, guiding the lympho-
activation energy Difference in free energy between the ground- cyte into the appropriate tissue.
state energy of a substrate and the transition state for the adduct (1) A type of DNA damage in which a foreign chemical
conversion of substrate into product. group is inappropriately attached to the DNA molecule. (2) A
activator A regulatory molecule, usually a protein, that increases product of a chemical addition reaction.
gene expression. adenine (A) Purine base that is one of the four types of bases in
active site The part of an enzyme to which substrate binds and RNA and DNA.
where conversion to product is catalysed.
adenosine deaminase deficiency (ADA) A deficiency of the
active transport Transport of molecules or ions across a
purine salvage enzyme adenosine deaminase, which results in
membrane against a concentration or electrochemical gradient
severe combined immunodeficiency (SCID) due to toxic effects
and which thus requires energy. on lymphocyte development.
active zone Region within an axon terminal where synaptic
adenosinetriphosphatase See ATPase.
vesicles fuse with the plasma membrane to release neurotrans-
mitter into the synaptic cleft. Proteins involved in the docking and adenosine triphosphate (ATP) A small, water-soluble nucleotide
fusion of synaptic vesicles are concentrated at this site. that is the principal carrier of energy in cellular metabolism. It is
activism Confronting and attempting to improve problematic formed by the phosphorylation of adenosine diphosphate (ADP)
situations. during the light reaction of photosynthesis and during respira-
activity coefficient Ratio of the activity of a solute to its tion.
actual molar concentration. It is the fraction of the actual adhesins Proteins on the surface of bacteria that mediate adhesion
concentration that appears to be present as measured by some to host cells by binding to receptors on the host cell.
physical process. adhesion molecules (1) Cell-surface proteins of various types
acylenzyme Enzyme to which an acyl group is covalently bonded through which cells bind to other cells or to the extracellular
by the reaction between a nucleophile on the enzyme and an matrix. (2) Proteins produced by granulocytes in some inverte-
acylating agent. brates. They are released by exocytosis and may act as opsonins
ADA See adenine deaminase deficiency. for infecting antigens. Adhesion molecules are also located on the
adaptation (1) A feature of an organism whose presence can be plasma membrane.
explained by the fact that it served some fitness-enhancing adhesion receptor A cell-surface protein that confers mechanical
function in the organism’s ancestors, allowing those organisms stability on cell–cell or cell–matrix interactions by binding to
that possessed the trait in question to out-reproduce those that particular ligands. Also known as adhesion molecule.
did not. (2) Changes in the function or behaviour of an organism adipohaemocyte A rarely seen type of haemocyte in inverte-
made in response to changes in the external environment. brates, characterized by refringent intracytoplasmic lipid
adaptive Describes a feature or behaviour of an organism that droplets.
serves to enhance that organism’s fitness. adjuvant Any substance that when mixed with an antigen
adaptive immunity The antigen-specific immune responses nonspecifically enhances the immune response to that antigen.
mounted by B and T lymphocytes, which give rise to antibodies, admixed population A population originating from the mixing of
effector T cells and immunological memory specific for the individuals from two or more ancestral populations, but having
antigen that evoked them. This type of immune response can be an identity of its own distinguishable from the parental
mounted against an almost infinite variety of antigens and is populations.
found only in vertebrates. Subsequent re-exposure to the antigen adoptive transfer The transfer of immune system cells or cell
produces a more rapid and stronger immune response, due to the derivatives into an animal or human to initiate or augment an
presence of immunological memory. immune response.
adaptive landscape A graphical representation of all possible
adrenal cortex The outer layers of the adrenal gland, which are
frequencies of a given set of alleles in a population, which confer
responsible for synthesis and secretion of corticosteroid hor-
different degrees of fitness. ‘Peaks’ and ‘valleys’ in the landscape
mones (e.g. cortisol, aldosterone).
represent combinations of allele frequencies of high and low
adrenaline (epinephrine) The main catecholamine hormone
fitness, respectively.
released by the adrenal medulla.
adaptive radiation Rapid evolutionary divergence of a single
phyletic line, e.g. the mammals, into different ecological niches adrenal medulla The inner core of the adrenal gland. It is made up
or adaptive zones, with the formation of many different groups. of chromaffin cells, which are developmentally related to
The different groups arise almost synchronously, so that in the adrenergic neurons. They synthesize and secrete the catechola-
fossil record they are perceived as originating at about the same mine hormones – noradrenaline and adrenaline.
time. adventitia External connective tissue layer of a blood vessel wall.
adaxial Facing towards the main growth axis. For example, the adventitious root Root formed on tissue other than young root
upper surface of a leaf is the adaxial surface, as it faces towards the tissue (apart from the embryonic root).
direction of growth of the stem. AER See apical ectodermal ridge.

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Glossary

aerobic metabolism The biochemical processes that occur when algaean An outer cell wall component of some chlorococcalean
an organism is breaking down glucose in the presence of oxygen green algae that consists of long polymethylenic chains associated
(aerobic respiration). with amide groups and minor amounts of N-alkyl-substituted
aerobic respiration The breakdown of glucose by cells in the pyrroles.
presence of molecular oxygen. alginate A polysaccharide composed of D-mannuronic acid and
aeropyles External openings in the chambered chorion of tardi- L-glucuronic acid residues in irregular sequences, found primarily
grade eggs and in many insect eggs, which permit gas exchange in some brown algae. Alginate is used as a thickener and
with minimal water loss. stabilizing agent in the food and pharmaceutical industries.
aetiological Causative. algorithm A set of rules for solving a problem in a finite number of
aetiological mutation Allelic variants directly predisposing to a steps.
disease. alkaline enzymes Enzymes whose activities are optimal at alkaline
aetiology The underlying cause of a disease or other pathological pH (410). Alkaliphilic microorganisms produce various extra-
condition. cellular alkaline enzymes, e.g. cellulases, amylases and proteases.
afebrile Without fever. alkaliphiles Microorganisms that grow optimally or very well at
afferent Carrying towards. pH 4 9, often between pH 10 and 12, but which cannot grow, or
afferent fibre A nerve fibre that is derived from a sensory grow only slowly, at neutral pH.
neuron and connects a peripheral sensory structure to the alkaptonuria A mild recessive genetic disorder of humans in which
brain. the metabolic breakdown of tyrosine is abnormal because the
enzyme homogentisate oxidase is missing. Affected individuals
afferent nerve fibres Nerve fibres that conduct from the periphery
accumulate homogentisic acid, which causes their urine and
to the central nervous system. The term broadly corresponds to
cartilaginous parts of their bodies to blacken.
sensory fibres.
alkylating agents Reactive chemicals that can transfer alkyl
afferent projection The axons that project from a structure to its
groups to cellular macromolecules, including DNA, RNA and
target, providing input to the target.
proteins.
affinity The binding strength of one molecule for another,
allele One of two or more alternative forms of a gene that can
considered as the strength of binding of a single binding site to
occupy the same position, or gene locus, on the chromosome. For
a monovalent ligand, e.g. ligand to receptor, substrate to enzyme
some genes, one allele is present in the great majority of the
or a single antigen-binding site on an antibody to its correspond-
population and is considered the normal or ‘wild-type’ form. For
ing antigenic determinant. cf. Avidity
other genes, there are numerous different alleles within the
afimbrial adhesins Bacterial adhesins that are not associated with
population, none of which clearly predominates.
fimbrial structures.
allele frequency For a given gene in a given population, the
afterhyperpolarizing potential A brief membrane potential frequency of occurrence of a particular allele, expressed as a
more negative than the resting potential that occurs after an fraction of all the alleles.
action potential. It is due to the conductances through voltage- allelic heterogeneity Phenotypic differences or similarities pro-
dependent potassium channels (and calcium-activated potassium duced by different mutations at a single gene locus.
channels in some neurons) that open as a result of the action allergen An environmental protein antigen or small-molecule
potential. hapten that provokes an allergic reaction in a sensitized individual.
agammaglobulinaemia The absence of immunoglobulins, and On initial encounter (sensitization) the allergen induces the
hence of antibodies. production of allergen-specific IgE antibodies. Subsequent en-
aganglionosis The absence of parasympathetic ganglion cells in counter with the allergen induces allergic reactions mediated by
the colon. these antibodies. Common allergens are the proteins in pollens and
agar A sulphated polysaccharide rich in galactose- and 3,6- other substances that provoke hay fever and asthma.
anhydrogalactose that is found in the walls of some red algae allergy An adverse immune reaction against a common environ-
and used in the food, medicinal and dental industries. mental antigen (the allergen). It occurs only in certain individuals.
agarose gel electrophoresis Electrophoresis carried out in an alloantigen An antigen found only in some members of a species.
agarose gel, which is used to separate DNA fragments between allocation of carbon The flow of assimilated carbon into various
100 bp and 50 kb in length. compartments or biochemical pathways.
age rank In a clade of N taxa, each taxon has an age rank from 1 to allogeneic Describes tissues, cells or molecules from one individual
N reflecting the order of first appearances, with the oldest given that provoke an immune reaction in another individual of the same
the rank of 1 and the youngest a rank of N. species. Allogeneic reactions are due to the genetic differences
agonist A drug or natural ligand that binds to and activates a between individuals, which are reflected in antigenic differences in
receptor. some of the proteins and other macromolecules they make.
AIDS Acquired immune deficiency syndrome. A severe immuno- allogeneic mouse strains Strains of mice that possess different
deficiency due to the depletion of T lymphocytes caused by major histocompatibility genes, and thus produce different major
infection with the human immunodeficiency virus (HIV). histocompatibility antigens.
airway hyperresponsiveness An exaggerated bronchoconstric- allograft A graft of tissue to a genetically nonidentical individual of
tor response to various stimuli, which results in bronchospasm the same species. Allografts are usually rejected, due to the
and increased obstruction of the airways. differences in major and minor histocompatibility antigens
algae General name for several taxonomic groups of single-celled between unrelated individuals. Also known as a homograft.
or simple multicellular photosynthetic eukaryotic organisms, alloimmune Describes the immune response to a genetically
including the seaweeds. nonidentical graft from an individual of the same species.

145
Glossary

alloimmunization An immune response mounted by an individual amastigote The form of the parasitic protozoan Leishmania in the
to ‘foreign’ antigens from another member of the same species. mammalian stage of infection.
Examples of antigens that produce alloimmunization are MHC amber Fossilized resin, usually from coniferous trees.
molecules and blood group antigens. ambisexuality Occurrence of male and female function in a single
allometry (1) Effects of differences in size on the structure and individual, either as simultaneous or sequential hermaphrodit-
function of organs and organisms. (2) The study of the relationship ism.
between size or mass, and the shape, structure and function of an amino acid Organic acid of the general formula
organism or part of an organism. RCH(NH2)COOH (a amino acid), where R is a distinctive side
allomorph One of the crystalline forms of a chemical compound. chain. They are the building blocks of proteins and precursors of
alloparapatric species A species that is thought to have arisen by a many other organic molecules. The 20 amino acids specified by
two-stage model of speciation in which allopatric speciation is the genetic code are all L isomers.
followed by completion of speciation by parapatric hybridiza- aminoacyl-adenylate Mixed anhydride formed between the a-
tion. carboxylate of an amino acid and the a-phosphate of adenylic
allopatric Occupying separate geographic areas. acid (AMP).
allopatric speciation Origin of new species through geographic
aminoacylation Reaction by which an amino acid is attached to
isolation of two distinct populations.
the 31 end of its appropriate tRNA. Also known as charging of the
allopatry Geographic isolation of two species or populations.
tRNA.
allopolyploid A polyploid that originated by the doubling of the
chromosomes of a zygote with two unlike chromosome sets, aminoacyl-tRNA Transfer RNA (tRNA) carrying the appropriate
0

usually as a result of the hybridization of two species. amino acid esterified to the 3-OH of the 3 -terminal adenosine
allosteric Describes an enzyme or other protein that can adopt at residue of the tRNA.
least two different tertiary structures that differ in their activity. aminoacyl-tRNA synthetases Enzymes that specifically attach
allosteric effect Change in protein activity produced by the amino acids to their appropriate tRNAs (aminoacylation).
binding of another molecule to a site outside the active site. aminoglycosides A family of antibiotics (mostly naturally occur-
allosteric effector A molecule that can activate (or inhibit) an ring products of Streptomyces or Micromonospora species) that
enzyme or other protein by binding to a site (the allosteric site) inhibit bacterial protein synthesis. Members of the group include
distinct from the active site, and inducing a change in the protein’s streptomycin, gentamicin, tobramycin and amikacin.
conformation. Such effectors can be structurally quite different ammonite Member of an extinct group of the molluscan class
from the substrate of the enzyme. Cephalopoda, having a chambered shell.
allosteric proteins See allosteric. amniocentesis Aspiration of the amniotic fluid that surrounds the
allostery The long-range interaction that can occur between fetus, in order to diagnose abnormalities.
spatially distant ligand-binding sites on certain protein mole- amniote Vertebrate with an embryo that is surrounded by an
cules, which is due to a change in protein conformation induced amnion. Reptiles, birds and mammals are amniotes
by binding of ligand at one of the sites. amniotic egg An enclosed egg with a shell and amniotic
alopecia Hair loss. membranes that protect and nourish the developing embryo.
alpha-satellite Repetitive DNA sequences present in great abun- Laid by reptiles, birds and egg-laying mammals. Also called a
dance in and around the centromeres of all the human cleidoic egg.
chromosomes. amoebal-plasmodial transition Differentiation of the CL strain
alternation of generations (1) Life history characterized by the of amoebae into a multinucleate plasmodium without mating.
presence of two phases, one haploid (the gametophyte) and one amoebocyte An immunocyte (haemocyte) of some invertebrates,
diploid (the sporophyte). (2) Sequential development of sexual synonymous with the archaeocyte of Porifera and coelenterates
medusoid stages from asexual polyp in jellyfish. and the granulocyte of Annelida, Mollusca, some Arthropoda and
alternative splicing The process whereby different proteins can be Echinodermata.
generated from a single gene by the choice of different splicing
AMPA receptors A group of receptors that respond to the
sites during the processing of the RNA transcript of the gene.
excitatory neurotransmitter glutamate and are distinguished
altricial Describes young that are relatively underdeveloped and
from other glutamate receptors by their selective response
helpless at birth, with limited sensory abilities and movement.
to a-amino-3-hydroxy-5-methyl-4-isoxazole proprionic acid
Some features needed for survival may be more advanced.
(AMPA).
alvarezsaurids Group of flightless, stout-forelimbed birds from
the Late Cretaceous that includes the Asian Mononykus amphimixis Type of sexual reproduction in which eggs and sperm
olecranus, Shuvuuia deserti and Parvicursor remotus, and the come from separate individuals.
South American Patagonykus puertai and Alvarezsaurus calvoi. amphipathic Describes a molecule in which one end is hydro-
alveoli (1) Flattened fluid-filled vesicles typically forming a layer phobic and the other is hydrophilic.
beneath the surface membrane of some protozoa. They form an amphiphile Surface-active molecule with a hydrophilic group at
integral part of the complex outer layer of cells in ciliates. (2) one end and a hydrophobic chain at the other.
Plural of alveolus. amphiphilic Having both water-soluble and water-insoluble
alveolus Individual terminal air sac in the lung. The human lung regions.
contains approximately 300  106 alveoli. amphisbaenians A group of long-bodied, generally limbless
Alzheimer disease A human degenerative disorder characterized squamate reptiles. Their body scales are arranged into rings
by progressive dementia. (annuli) and they superficially resemble stout earthworms. One
amantadine An inhibitor of the Influenza A virus. genus retains front limbs.

146
Glossary

amphotericin B A member of the polyene class of antifungal aneuploid, aneuploidy Having too many or too few chromo-
drugs. It has been used for over 30 years to treat a variety of somes, or chromosome segments, compared to the normal
serious invasive fungal infections. genotype. Usually refers to the presence of an extra copy of a
ampullae of Lorenzini Electroreceptor organs located on the single chromosome (trisomy) or the absence of a single
heads of sharks. chromosome (monosomy).
amyloid A proteinaceous deposit arising from incorrectly folded, aneurogenic limbs Limbs of larval Ambystoma that develop in the
protease-resistant peptides. It is present in pathologies such as absence of innervation.
neurofilament tangles in Alzheimer disease. aneurysmal Abnormally enlarged or dilated.
amyloidosis Extracellular deposition of an insoluble protein com- anf Gene designation for the iron-only nitrogenase system.
plex, usually derived from serum proteins, with a fibrillar structure angiogenesis Formation of new blood vessels.
and a characteristic twisted b-pleated sheet conformation. angioplasty A process whereby restenosis of an artery following
amylolytic Describes starch degradation by hydrolytic enzymes, surgery is prevented by insertion of a balloon device within the
that is, by amylases. artery.
amyloplast A form of mature plastid filled with starch grains and angiosperm A plant with seeds that are produced in an ovary.
found in plant storage tissues. Angiosperms are also known as the flowering plants.
anabolism Constructive metabolic reactions in which complex 3,6-anhydro ring An internal, cyclic ether formed by formally
molecules are synthesized from simpler ones and tissues built up. eliminating HOH between the hydroxyl groups at positions C3
anaemia Abnormal condition in which there is a reduced number and C6.
of red blood cells per unit volume of blood. This reduction may animal cap In amphibian embryos, the area of smaller, more
also be expressed as decreased concentration of haemoglobin in rapidly dividing cells around the non-yolky animal pole.
the blood, or decreased packed cell volume (haematocrit). animal venom Complex mixture of substances produced by an
anaerobic Occurring in the absence of oxygen. animal in which one or more components are toxins.
anaerobic glycolysis Metabolic pathway for the breakdown of anlage (plural anlagen) The primordial undifferentiated stage of
glucose in the absence of oxygen. a structure in an embryo or larva.
anaerobic metabolism The biochemical processes that occur Annelida, annelids The ringed or segmented worms. A large
when an organism is breaking down glucose or other respiratory grouping of mainly marine animals that also contains terrestrial
substrate in the absence of oxygen (anaerobic respiration). groups such as earthworms and leeches.
anaerobic polypeptides (ANPs) A small set of proteins that are
anomer One of the two isomers (a or b) of a monosaccharide as
synthesized in roots and shoots of young seedlings in an
determined by the stereochemistry at the carbonyl carbon atom.
anaerobic environment.
Anopheles A genus of mosquitoes that includes species capable of
anaerobic respiration The breakdown of glucose or other
transmitting malaria parasites to mammals.
respiratory substrate by cells in the absence of molecular oxygen.
anorexia Diminished appetite or aversion to food.
analogous Describes features, including proteins, that are similar
anoxia The complete absence of oxygen.
by convergent evolution.
analogy Similarity between characters or character states that is anoxygenic photosynthesis Type of photosynthesis in which
oxygen is not generated.
not due to a common origin but is generally explained through
independent adaptation. ANPs See anaerobic polypeptides.
anamorph An asexual, usually haploid yeast culture, of a single antagonistic interaction (1) An interaction between two species
mating type. that results in one of the species increasing its fitness while the
anaphylatoxins Complement peptides C3a, C4a and C5a, which fitness of the other is reduced. (2) An interaction of a drug with a
are powerful systemic triggers of inflammation and can thus receptor that inhibits the usual function of the receptor.
induce systemic anaphylactic shock. antennae The complexes of light-harvesting molecules in photo-
anaphylaxis A rapid life-threatening systemic allergic reaction that synthetic systems.
can result when a sensitized individual encounters an allergen. It anterior Towards the ‘head’ end of an animal.
is characterized by tracheal swelling and systemic shock resulting anthelmintic A drug used to eradicate intestinal parasites.
in complete circulatory system shutdown. anther The organ in a flower in which the pollen, which contains
anaplerotic Associated with biosynthetic biochemical pathways. the male gametophytes, is formed.
anaplerotic reaction A reaction which replaces a component that anthocyanidin Red, blue or purple pigments with the basic
has been removed, usually from a cyclic reaction sequence, for use structure 2-phenylbenzopyrylium (or flavylium) ion, consisting
in biosynthesis. of two benzene rings (A and B) joined by an O-heterocyclic ring.
anapsid The primitive condition of the vertebrate skull in which An anthocyanidin combined with one or more sugars forms an
there are no temporal openings. anthocyanin.
andesite Volcanic lava of intermediate composition in the calc- anthropogenic Caused by human activities.
alkaline volcanic suite of rocks. anthropoid A general term of primate classification that includes
androgen Class of steroid hormones, including the male sex only monkeys, apes and humans.
hormone testosterone, produced by the gonads and adrenal glands. antibiogram The spectrum of susceptibility or resistance of a
anergy (1) A state of nonresponsiveness to antigen. Lymphocytes bacterial strain to multiple different antibiotics.
are said to be anergic when they cannot respond to their specific antibiotic peptides Small 2–7 kDa peptides that kill certain
antigen under optimal conditions of stimulation. (2) A lack of microorganisms. In most cases, they bind to phospholipids in
delayed-type hypersensitivity response to antigens to which there the cell membrane, associating and forming pores in the lipid
is universal exposure. bilayer.

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Glossary

antibodies Proteins secreted by the immune system in response antigenicity (1) Degree to which an antigen stimulates an immune
to an infection or to immunization with a foreign antigen. response. In this sense the term is synonymous with immuno-
Antibodies, or immunoglobulins, are highly variable proteins, genicity. (2) Degree to which an antigen reacts with a particular
specific to the antigen that evoked them. They bind to that antigen receptor.
antigen and aid its destruction and clearance from the body. They antigen-presenting cell (APC) A cell displaying antigenic peptide
are made only by the B lymphocytes of the immune system. An on its surface in a complex with MHC class I or class II molecules,
immunoglobulin molecule consists of two heavy (H) and two the form of antigen that can be recognized by T cells. The term
light (L) chains each containing one variable (V) region, which usually refers to macrophages, dendritic cells and B cells, which
forms the antigen-binding site, and one (L chains) or two to three also produce co-stimulatory molecules such as the B7 molecules,
(H chains) constant (C) regions. and are thus able to activate naive T cells specific for the antigen
antibody-dependent cell-mediated cytotoxicity (ADCC) Cell presented.
killing by natural killer (NK) cells of cells coated with antibodies antigen-specific cells T lymphocytes and B lymphocytes that have
specific for antigens on the cell surface. receptors specific for a particular antigen on their surface.
anticholinesterase Enzyme that breaks down acetylcholine at antiglobulin test A means of detecting antibody or complement
synapses, thus limiting its effect after a nerve impulse is passed on. fragments on red blood cells taken directly from the patient
This prevents the target nerve or muscle cell from being (direct antiglobulin test), or antibody to red blood cells in plasma
continually stimulated. (indirect antiglobulin test).
anticipation In genetics, a phenomenon characterized by the earlier antihypertensive Able to reduce high blood pressure, applied to
onset of an inherited disease in each succeeding generation. The drugs.
disease is also usually more aggressive in the newest generation. anti-idiotope A binding site within the antigen-binding site of an
anticipatory immunity See adaptive immunity. immunoglobulin or T-cell receptor that is specific for a particular
anticodon The three consecutive ribonucleotides in a transfer idiotope on a particular immunoglobulin.
RNA (tRNA) molecule that base pair directly with a comple- anti-idiotypic antibody An antibody that is directed specifically
mentary codon of three nucleotides in the messenger RNA against the idiotype unique to another antibody molecule.
(mRNA), thus enabling the genetic code to be translated into a anti-inflammatory agents Drugs that block or inhibit all or part
sequence of amino acids. of the inflammatory reaction.
antidromic Describes the passage of nerve impulses from the axon antioxidant Any substance that significantly delays or prevents
terminal of a neuron to its cell body. rather than in the more usual oxidation of an oxidizable substrate when present at a low
(orthodromic) direction – cell body to axon terminal. concentration compared with that of the substrate.
antifolate drug Substance that interferes with the synthesis of anti-peptide antibodies In biotechnology, antibodies elicited
folate and its derivatives, thereby blocking replication of DNA. against synthetic peptides.
antigen Any substance that can be specifically recognized by T and anti-phase Describes two oscillatory systems in which the time
B lymphocytes and elicit an adaptive immune response. Most between some measured event in each of them is exactly half a cycle.
antigens are proteins, but carbohydrates and nucleic acids can be anti-phosphoprotein antibodies Antibodies generated against
antigenic. Small molecules can act as antigens if attached to a particular phosphorylation sites of a phosphoprotein that can be
protein carrier. If an antigen is capable of inducing an immune used to distinguish between phosphorylated and non-phosphoryl-
response on its own, it is also called an immunogen. ated forms of the protein.
antigen-binding site The part of an immunoglobulin molecule antiporter A transmembrane protein that couples the transport of
that binds antigen. An antigen-binding site is formed by the two solutes across a membrane in opposite directions.
paired variable (V) regions of a heavy and a light chain. Each antisense RNA (1) Any RNA complementary to an mRNA and
immunoglobulin monomer thus contains two identical antigen- which is able to base pair with the mRNA and block its function.
binding sites. (2) In bacteria, a naturally occurring short, untranslated RNA
antigen presentation The display of antigenic peptides, derived transcript, often acting as a repressor of plasmid replication. The
from processed protein antigens, complexed with major histo- antisense RNA exerts its effect by binding to a complementary
compatibility complex (MHC) molecules on the surface of a cell. transcript, and modifying its secondary structure or preventing its
The complex of peptide and MHC molecule is the form of antigen translation.
recognized by T cells. antisense RNA injection Technique for analysing gene function in
antigenic determinant See epitope. Caenorhabditis elegans in which antisense RNA for a particular
antigenic drift Stepwise changes in the antigenic properties of a gene is injected into the gonad of a hermaphrodite worm, and the
protein caused by successive point mutations in the gene for the progeny studied for abnormal phenotypes.
protein. antisperm antibodies Immunoglobulins that react with sperm.
antigenic shift A major change in the antigenic properties of antitermination Mechanism that suppresses intrinsic transcrip-
certain viral protein antigens (e.g. haemagglutinin and neurami- tion termination signals through modification of RNA polymer-
nidase in influenza virus) that occurs when the RNA segments of ase activity.
a human strain of the virus reassort or exchange with those from antivenin A substance that can block, neutralize or reverse the
an animal strain of the virus. toxic effects of a venom.
antigenic site See antigenic determinant. antivenom See antivenin.
antigenic variation The ability of certain pathogenic microorgan- aperture In regard to computed tomography, the hole in the gantry
isms, such as trypanosomes and some bacteria, to vary their through which the patient is moved as the examination
surface antigens from generation to generation and thus evade the progresses. Ideally, this should be as large as possible, thereby
immune defences of the host. diminishing any sense of claustrophobia.

148
Glossary

apical complex Organelles (rhoptries, micronemes) at the apical arabinoxylan A polysaccharide composed primarily of L-arabino-
end of the invasive stages (merozoite, sporozoite, ookinete) of the syl and D-xylosyl units. In every case, the xylosyl units form the
malaria parasite. backbone structure, which may be branched or unbranched, with
apical ectodermal cap Epidermis covering the distal end of a the L-arabinosyl units present as single side units or in short
developing limb in amphibian embryos. It is homologous to the oligomeric chains.
apical ectodermal ridge of chick embryos. arachidonic acid An unsaturated 20-carbon fatty acid that is the
apical ectodermal ridge (AER) Thickened ridge of ectoderm at major precursor for prostaglandin and leukotriene (eicosanoids)
the distal tip of the limb buds in avian and mammalian embryos biosynthesis.
that is necessary for proximo-distal growth of the limb. It arachnoid villi Microscopically visible outpouchings of the arach-
produces fibroblast growth factor signals. noid through the dura mater, protruding into the lumen of venous
apicomplexans Protozoan parasites of the phylum Apicomplexa sinuses.
which are characterized by a set of subcellular organelles located arboreal (1) Living in trees. (2) Adapted for life in the trees.
at the apical end of the parasite that have important roles in host- arborescent (1) Describes a plant architecture resembling that of a
cell invasion. tree but in which the trunk is not constructed primarily of
apiculate Describes a lemon-shaped yeast cell. secondary wood. (2) Branched like a tree, applied e.g. to dendrites
aplacental yolk sac viviparity A form of uterine incubation of a of neurons.
yolk-dependent embryo in which the uterus does not augment arboviruses Arthropod-borne viruses, viruses transmitted by
yolk stores to provide additional nutrients. insects and other arthropod vectors.
apodemes Downward-projecting knobs on the underside of the arbuscule The much branched ‘tree-like’ hyphal structure pro-
arthropod exoskeleton, for the attachment of ligaments or duced inside plant cells by certain mycorrhizal fungi (arbuscular
muscles, and limb bases. mucorrhizal fungi). The major processes of nutrient exchange
apolipoprotein The protein component of the complexes of lipid between the partners are thought to occur across the membranes
and protein carried in blood plasma. investing these branches.
Archaea A kingdom of prokaryotic organisms that forms one of
apomorphic Describes an evolutionarily novel character that has
the two major divisions of the prokaryotic world. Archaea differ
appeared in the phylogenetic group in question (a derived
as much from other prokaryotes (the Bacteria or eubacteria) as
character), as opposed to primitive or plesiomorphous charac-
both do from eukaryotes. Many live in extreme conditions.
ters, which are characters inherited by that group.
archaeal virus, archaeal phage Virus that infects an archaeal
apomorphy See apomorphic.
host. Also known as an archaeophage.
apomixis Asexual reproduction without meiosis and fertilization
archaeocyte A motile amoeboid cell found in the mesopyl of
in plants, which can involve cells other than the ovule.
sponges. It is nucleolate, capable of phagocytosis and able to
apoplasm Cell walls and other compartments outside the cell develop into any other sponge cell type.
membranes. Archaeopterygids The most primitive group of birds, including
apoptosis A type of physiological cell death in multicellular the Late Jurassic Archaeopteryx lithographica from Germany and
animals, also known as programmed cell death, in which the Late Cretaceous Rahonavis ostromi from Madagascar.
an internal death programme is activated in the cell. This leads archetype An idealized image of the most fundamental ‘ground
to DNA fragmentation, nuclear condensation and surface plan’ upon which the structure of the group of animals or plants is
changes that result in the dying cell being rapidly phagocytosed based.
by macrophages. Apoptosis is essential in both embryonic Archosauria A group of amniote (laying shelled eggs) animals that
and adult tissues to eliminate unwanted or potentially includes not only the living crocodiles and birds, but also
harmful cells and maintain cellular homeostasis. It contrasts dinosaurs, flying reptiles (pterosaurs) and a range of more
with necrosis, which is a lytic form of cell death induced by toxic obscure Permian and Triassic amniotes.
agents. Ardipithecus A genus of primitive hominins established in 1995 by
aposymbiotic host A host whose symbionts have been removed. Tim White and colleagues. At present it contains only one species,
appetitive conditioning A classical conditioning protocol based A. ramidus.
on pairing a neutral conditioning stimulus (CS) with a rewarding areoles Spaces between the grooves that pass over the epicuticle of
unconditioned stimulus (US) such as food. arthropods.
AQP See aquaporin. aromatase Enzyme that catalyses the conversion of testosterone to
aquaporins Family of water-channel proteins present in the oestradiol (also androstenedione to oestrone).
plasma membrane (plasma membrane intrinsic protein, PIP) ARP complex A complex of proteins, including two actin-related
and tonoplast (tonoplast intrinsic protein, TIP). They are proteins, that is implicated in controlling actin polymerization.
identified by their permeability to water and by the signature arthroconidium A conidium that results from fragmentation of a
Asn-Pro-Ala sequence present in both the amino- and carboxy- septate hypha. Also called an arthrospore.
terminal halves of the molecule. Arthus reaction, Arthus response Typical type III hypersensi-
Aquifex pyrophilus Hyperthermophilic rod-shaped bacterium that tivity reaction that is induced a few hours after injecting a soluble
forms water during growth on hydrogen and oxygen. antigen into the skin of a sensitized animal. The formation of
arabinogalactan A polysaccharide composed primarily of L- immune complexes causes an inflammatory reaction at the site of
arabinosyl and D-galactosyl units. In every case, the galactosyl antigen injection that is characterized by local oedema, haemor-
units form the backbone structure, which may be branched or rhage and necrosis.
unbranched, with the L-arabinosyl units present as single side artiodactyl mammals Hoofed mammals with an even number of
units or in short oligomeric chains. functional toes on each foot, such as camels, bison and cattle.

149
Glossary

AS-C See Achaete-scute complex. atrium (1) Concave ventral side of a zooid, surrounded by the
ascites Collection of fluid in the peritoneal cavity. In the context of tentacles. (2) Upper thin-walled chamber of the vertebrate heart.
haemolytic disease of the newborn, ascites is due to congestive atrophy A wasting away or diminution in the size of a cell, tissue,
cardiac failure. organ or part.
ascocarp A complex fruiting body containing sac-like asci attachment In a viral infection, the binding of a virus particle to a
produced by ascomycete fungi. specific receptor on the surface of a host cell.
ascomycete A member of the ‘sac’ or ‘cup fungi’, phylum attenuated strain A non-pathogenic form of a normally patho-
Ascomycotina. They are characterized by the production of genic microorganism. It retains the capacity to evoke an immune
sexual spores within a sac-like structure, the ascus, the sacs response but lacks the capacity to do harm within the host. Such
sometimes being borne in cup-shaped fruiting bodies. strains can arise naturally or be produced by prolonged culture in
ascus The sexual spore-bearing structure in ascomycete fungi, the laboratory.
which carries the meiotic products (ascospores). attenuated vaccine A vaccine consisting of a living attenuated
aseptic necrosis Tissue death occurring in the absence of infection. strain of the relevant microorganism. Such strains are unable to
aseptic technique A set of working practices designed to minimize cause disease but can still cause a limited infection in the host and
the risk of biological contamination of the work. induce immunity.
asexual proliferation Division of a cell without the exchange of attenuation (1) Regulated arrest or termination of transcription
genetic material. due to structural alterations of transcriptional pausing or
asexual reproduction Reproduction that does not involve meiosis termination sites. (2) See attenuated strain.
and fusion of gametes, and which produces progeny genetically attractor An object in phase space to which the trajectory
identical to the parent. ultimately comes (and stays) arbitrarily close.
aspergillosis Disease caused by invasive infection by fungi of the auditory bulla The bony capsule surrounding the sensory struc-
genus Aspergillus. tures of the ear.
asplenia The total absence of a spleen. aufwuchs community All the sessile microorganisms living on any
assembly The stage of viral replication during which all the animate or inanimate substrate. The term is mostly used in the
structural components of the virus come together at one site in the context of aquatic microorganisms.
cell to form the basic virus particle. AUG Adenine-uracil-guanine, the methionine codon used as the
association by colonization The acquisition of a parasite start signal for translation.
from another host that is not itself ancestral to the present augmentation In short-term synaptic plasticity, an Intermediate
host. phase of enhanced transmission lasting several seconds.
association by descent The inheritance of a parasite from a host aura Warning symptoms that occur before the headache in about
that is ancestral to the present host. 25% of migraine sufferers. It is most often visual, but weakness or
assortative mating A tendency for the characteristics of mating numbness of one side of the body and speech disturbance may
partners to be correlated. For example, large males may tend to occur.
mate with large females (positive assortative mating) or pale- AURE Adenosine uridine-rich element. A sequence motif confer-
coloured males with dark-coloured females (negative assortative ring instability on mRNA. It is located in the 30 -untranslated
mating). region of some mRNAs in mammalian cells. Different AURE-
astragalus The bone in the ankle which has a joint with the tibia binding proteins (AUBPs) have been identified.
(shin bone) proximally and the navicular and cuboid (other ankle australopithecine Informal name describing the species included
bones) distally. in the hominin subfamily Australopithecinae. The subfamily
astrocyte Non-nervous support cell type that constitutes the includes the genera Ardipithecus, Australopithecus and
‘connective tissue’ of the central nervous system. Paranthropus.
ataxia telangiectasia Complex multisystem disorder character- Australopithecus The genus name introduced by Raymond Dart in
ized by neurological impairment, variable immunodeficiency and 1925 for the first discoveries of early hominins in Africa.
dilation of ocular and cutaneous capillaries. autacoids Substances produced in physiological or pathophysio-
atomic force microscope A microscope that scans a fine tip logical responses to injury. They have a brief lifetime and
(radius 1–20 nm) over a surface and provides images of solids, generally act locally, near the site of synthesis. Examples are
including biological structures, with atomic accuracy (lateral histamine, bradykinin, and 5-hydroxytryptamine.
resolution of 1 nm, vertical resolution 0.1 nm). The image is autapomorphy A derived character, a unique character possessed
formed by the continuous measurement of forces between the tip by a taxon.
and sample. auto A prefix denoting derived from self or acting on self. See
ATP See adenosine triphosphate. autoantigen, autoantibody, autoimmune disease.
ATPase Adenosinetriphosphatase. Enzyme activity that hydro- autoantibodies Antibodies produced by an individual to consti-
lyses ATP to give ADP and inorganic phosphate. tuents of their own tissue.
ATP-dependent H1 pump See H1 ATPase. autoantigen A constituent of an individual’s own tissues to which
ATP synthase A membrane-bound protein complex that synthe- the immune system mounts an immune response.
sizes ATP from ADP. It uses the energy stored in the proton autoclave A pressure vessel used to sterilize materials by use of
electrochemical gradients across the thylakoid membrane of compressed steam, typically at 0.1 to 0.2 MPa (15 to 30 psig) and a
chloroplasts or the inner mitochondrial membrane. high temperature.
atresia A congenital complete closure of a normal body orifice or autocrine Describes the action of a hormone, cytokine or other
organ, such as the oesophagus. secreted factor when it acts on the cell that has produced it.

150
Glossary

autocrine growth factor A secreted polypeptide that acts on the autosomal set One copy of each type of non-sex chromosome in an
same cell that produced it, stimulating proliferation or differ- organism. A normal diploid female mammal, for example, is
entiation. 2X:2A, meaning that her cells contain two X chromosomes and
autograft A transplant of tissue from one site to another in the two autosomal sets (two copies of each autosome).
same individual. autosome Any chromosome other than the X and Y sex
autoimmune disease Any disease in which the immune system chromosomes.
attacks the body’s own cells. Disorders in this category include autotoky Type of reproduction characterized by the production of
acquired haemolytic anaemia, rheumatic fever, insulin-depen- progeny by a single parent. It includes hermaphroditism and
dent diabetes and myasthenia gravis. parthenogenesis.
autoimmune haemolytic anaemia A condition in which a person autotroph, autotrophic organism An organism that uses carbon
produces antibodies to antigens on their own red blood cells, dioxide as the sole carbon source and can use inorganic sources for
leading to destruction of red cells. the other essential elements such as nitrogen. Autotrophs include
autoimmune uveitis A degenerative eye disease characterized by the plants and photosynthetic bacteria, which use sunlight as the
inflammation of the uvea (iris, ciliary body and choroid) which is energy source, and chemoautotrophic bacteria, which use inorganic
caused by the generation of antibodies against antigens in these chemicals as energy sources. Directly or indirectly, autotrophs
tissues. support all the other organisms in an ecosystem.
autoimmunity Immune responses to components of the body’s autotrophic See autotroph.
own tissues, which can result in tissue damage and inflammation auxotroph A genetic variant, usually of a microorganism, that has
leading to autoimmune disease. nutritional requirements in addition to those of the wild type. In
autolysis Cell lysis catalysed by the cell’s own enzymes. this context the latter is known as a prototroph.
autonomic nervous system The parasympathetic and sympa- auxotrophy Having nutritional requirements in addition to those
thetic nervous systems, which are responsible for largely of the wild type.
unconscious regulation of visceral functions such as heartbeat, aversive conditioning A classical conditioning protocol based on
smooth muscle contraction and relaxation, and glandular pairing a neutral conditional stimulus (CS) with an aversive
secretion. unconditional stimulus (US) such as an electric shock.
autonomy The capacity to be self-determining. Autonomous avidity The summation of the individual binding strengths, or
agents are able to direct their lives by making choices in affinities, when molecules with multiple binding sites bind to each
accordance with some underlying value structure that the other. An example would be the overall strength of binding when
individual has. an antibody with multiple binding sites (e.g. IgM) binds to an
autophagic vacuoles Membrane-bounded cellular organelles that antigen with multiple identical epitopes.
contain cytoplasmic contents but no lysosomal hydrolases.
avifauna In palaeontology, an assemblage of fossil birds from a
autophagosomes Membrane-bounded cellular organelles result-
single deposit.
ing from fusion of lysosomes and autophagic vacuoles, in which
avirulence factor (Avr factor) In plant pathology, a product of a
the digestion of contents occurs.
pathogen that specifically elicits expression of a dominant
autopolyploid A species in which the genome consists of multiple
resistance gene on a gene-for-gene basis.
identical copies of a basic set of chromosomes, arising from the
avirulence gene (Avr gene) In a plant pathogenic microbe, a gene
doubling of an original chromosome set.
that specifies an avirulence factor.
autoradiography Photographic process that captures the location
avirulent A form of an infectious agent that does not cause
of a radioisotope-labelled substance that has been used to probe an
significant disease.
in vivo or in vitro system. The radiation emitted by the labelled
molecule or structure is recorded on a photographic plate (light avulsion Tearing away or forcefully extruding a segment of bone or
microscopy) or on thin, self-supporting film (electron microscopy) other tissue.
autoregulation (1) Situation in which an enzyme acts to activate axillary Pertaining to, or situated near, the axilla, or armpit.
or inhibit its own activity. (2) Process by which a protein inhibits axis formation Formation of the main body axis, which occurs
further synthesis of itself. (3) Of blood flow, the ability of blood early in development. In vertebrate embryos, it refers to the
vessels to respond to chemical and physical signals in their local establishment of the prechordal plate and the notochord, the
environment, i.e. non-neuronal signals, and adjust blood flow to main components of the axial mesoderm.
capillary beds to meet the local demand. axolemma The membrane surrounding an individual neuron.
autosomal dominant A mutation or allele of an autosomal gene axon A long thin process (0.5–2.0 mm in diameter) originating
(a gene on any chromosome other than the X and Y sex from the cell body of a nerve cell (neuron) and specialized to
chromosomes) that determines the phenotype when only one transmit information. It conveys electrical impulses from the cell
copy is present in the diploid genome (the heterozygous state). body to the axon terminals, where the signal is passed on via
autosomal recessive A mutation or allele of an autosomal gene synapses to other neurons or target cells.
(a gene on any chromosome other than the X and Y sex axonal transport The active transport of cytoplasmic constituents
chromosomes) that only determines the phenotype when two within the axon. Transport occurs in both directions within the
copies are present in the diploid genome (the homozygous state). axon.
autosomal recessive inheritance The pattern of inheritance of a axoneme The core of any cell appendage based on microtubules,
trait that is determined by an autosomal recessive allele. Parents such as cilia, axopodia or suctorian tentacles.
heterozygous for the allele do not show the trait, but for each of axon guidance During the development of nervous systems, the
their offspring there is a 1 in 4 chance that they will be response of the highly mobile growth cone of migrating axons to
homozygous for the allele and thus display the trait. environmental cues that guide them to their targets.

151
Glossary

axopodia Long, stiff, tapering cytoplasmic projections from the bacteriolytic Able to cause sufficient damage to the bacterial cell
surface of certain protozoa. wall to cause lysis of the bacterial cell. Lysozyme and penicillin are
azole Type of antifungal agent introduced over the past 20 years, examples of bacteriolytic molecules.
which is used to treat a range of fungal infections. bacteriophage k cI protein A transcription factor encoded by the
azygospore A spore that is morphologically similar to a zygo- bacteriophage l genome that regulates phage gene expression
spore, but is of asexual origin. during lysogeny.
bacteriophage A virus that infects bacteria. Also known as phage.
bacteriostatic Able to stop bacterial growth, but not to kill,
leaving the bacteria able to resume growth and replication on
B cell Type of antigen-specific lymphocyte that produces
removal of the bacteriostatic agent
immunoglobulins (antibodies) and is responsible for humoral
immunity. It is produced in the bone marrow (or in the bursa of bacterium (plural bacteria) A member of the domain Bacteria.
Fabricius in birds) and develops independently of the thymus. baculovirus vector A DNA virus that infects insects and is often
After encounter with antigen, B cells differentiate into antibody- used as a vector to express foreign proteins in cultured cells.
secreting plasma cells. baculum A bone within the penis.
B DNA The most common form of double-stranded DNA helix in balanced genotype Having the normal number of each gene. A
biological systems. The helix is right-handed and the bases lie at cell with a reciprocal translocation has an abnormal karyotype
roughly 901 to the helix axis. The helix has distinct major and but retains a balanced genotype.
minor grooves. ballistoconidium Any spore that is forcibly discharged. Also
called a ballistospore,
B lymphocyte See B cell.
barbed end The fast-growing end of an actin microfilament.
B1 cells A subpopulation of B cells found in sites such as the
basal lamina A thin sheet of extracellular matrix, composed of
peritoneal cavity. Unlike the majority of B cells they are self-
proteoglycans and glycoproteins, particularly laminins, that lies
renewing and carry (in humans and mice) the cell-surface marker
immediately underneath an epithelium, or around muscle cells
protein CD5.
and Schwann cells, separating them from connective tissue. The
B2 (or conventional) B cells The vast majority of the B cells found basal lamina has important functions in controlling cell
in peripheral lymphoid organs. differentiation, cell polarity and cell division. Also known as
B800 Bulk bacteriochlorophyll a molecules with absorption basement membrane.
maximum at 800 nm. basal metabolic rate See basal rate of metabolism.
B850 Bulk bacteriochlorophyll a molecules with absorption basal plasma membrane Region of the plasma membrane of an
maximum at 850 nm. epithelial cell on the face of the cell away from the lumen.
BAC Bacterial artificial chromosome. A large-insert cloning system basal rate of metabolism The rate of metabolism in adult
that utilizes a small cloning vector based on the Escherichia coli F endotherms when they are at rest in a thermally neutral
factor. It incorporates an antibiotic-resistance marker, cloning environment after a short period of fasting. It represents the rate
sites and rare-cutter restriction sites for mapping. of energy expenditure for self-maintenance (homeostatic regula-
back focal plane The focal plane of a lens that lies behind it when tion) measured as oxygen consumption or production of heat or
viewed in the direction of illumination. This term is usually used carbon dioxide by resting animals.
when referring to the objective lens, where the phase ring, carried base (1) A substance that accepts protons from water or another
on the phase plate, is properly found. donor. A strong inorganic base is a compound that dissociates,
backprojection Method used to generate a three-dimensional producing hydroxyl (OH) ions. (2) Any of the nitrogen-contain-
structure from several different two-dimensional images of the ing ring compounds (purines or pyrimidines) that are constituents
object. of nucleotides. They are referred to as bases because they can
Bacteria One of the two domains of prokaryotes (the other is the combine with H 1 in acidic solutions. Also called a heterocycle.
Archaea), comprising most common microbes. base modification A change in the chemical structure and/or
composition of the purine and pyrimidine bases in RNA or DNA.
bacterial artificial chromosome See BAC.
base pair A pair of complementary nucleotides held together by
bacterial photosynthesis Type of photosynthesis found in some
hydrogen bonding in a double-stranded DNA molecule. In
bacteria, in which light harvesting by photosynthetic pigments is
DNA, adenine pairs with thymine and cytosine with guanine.
linked, via electron transport, to the generation of an electro-
base-exchange reaction An enzymatic reaction in which one
chemical proton gradient or reducing equivalent, but without the
class of glycerophospholipids is converted into another by
production of oxygen.
exchange of the polar head group.
bactericidal permeability-increasing protein (BPI) A cationic
basement membrane See basal lamina.
protein that binds and inactivates lipopolysaccharide in the outer
basic replicon The smallest piece of a plasmid which will replicate
membrane of Gram-negative bacteria, thus killing them.
to give the normal number of copies.
bactericidal Able to kill bacteria. basidiomycete Any fungus of the phylum Basidiomycotina. They
bacteriochlorophylls The bacterial equivalents of plant chloro- are characterized by production of sexual spores (basidiospores)
phylls. externally on cells known as basidia. The spores are often
bacteriocin A plasmid-encoded protein that is toxic to other produced in complex fruit bodies commonly known as mush-
bacteria. rooms and toadstools.
bacteriocyte Fat-body cells of aphids containing intracellular basidiospore A sexual spore in fungi of the phylum Basidio-
bacteria as symbionts. mycotina. It is borne on the outside of a basidium.

152
Glossary

basidium (plural basidia) A cell in fungi of the phylum b-glycosolic linkage Bond between C10 of the sugar ring and the
Basidiomycotina in which karyogamy and meiosis occur and base in a nucleoside or nucleotide, which is b because the base lies
which develops a definite number of basidiospores on its surface above the plane of the sugar.
following meiosis. b-lactam antibiotics Naturally occurring or semisynthetic anti-
basilar impression, basilar invagination Developmental process bacterial compounds characterized by a b-lactam ring that is
in which the floor of the posterior fossa of the skull is pushed essential for biological activity. Examples are penicillins,
superiorly by the bony elements of the upper portion of the cephalosporins, carbapenems and monobactams.
cervical spine. b-lactamases A family of bacterial enzymes that are able to
basolateral membrane The plasma membrane on the non- inactivate penicillins, cephalosporins or other b-lactam antibio-
luminal face of an epithelial cell. Nutrients are transported across tics by hydrolysis of the b-lactam ring.
the basolateral membrane of absorptive cells to the blood or b-pleated sheet A regular element of secondary structure in
intercellular space. proteins, in which two or more extended strands of the
basophil A type of polymorphonuclear white blood cell. It has polypeptide chain lie side by side (running either parallel or
receptors for IgE antibodies and is involved in allergic reactions. antiparallel), held together by a regular array of hydrogen bonds
It contains granules that stain with basic dyes. between backbone NH and C5O groups, to form a ridged planar
batholith Large magma chamber that formed at the base of a surface. The amino-acid side chains alternately face to opposite
volcanic chain and later cooled. sides of the sheet.
b-sheet structure See b-pleated sheet.
bauplan The basic body plan or body organization of an animal.
bilateral infiltrates Appearance of a localized, ill-defined opacity
Only about 30–40 different bauplans exist, and virtually all were
on both lungs in a chest radiograph, indicating airspace disease.
established by the early Cambrian period, 500–600 million years
Bilateria, bilaterians Animals with bilaterally symmetrical bodies,
ago.
or which are descended from phyla of bilaterally symmetrical
BBP Branch site-binding protein. See splicing factor 1. animals. All present-day bilaterians are triploblasts.
B-cell antigen receptor (BCR) See B-cell receptor bilayer A membrane or biomolecular leaflet consisting of two
B-cell receptor (BCR) The antigen-specific receptor on B cells. It monolayers of amphiphilic molecules.
consists of membrane-bound antigen-specific immunoglobulin bilayer defect A disruption in the regular bilayered structure of a
complexed with signal-transducing non-immunoglobulin Iga biological membrane that must occur for membrane fusion to
and Igb chains. Individual B cells carry B-cell receptors of a single proceed.
antigen-specificity. biliary contrast agents In radiography, contrast agents that are
B-cell tolerance The fact that the mature B cells present in an taken up by the liver.
individual do not normally respond to that individual’s own biliary Pertaining to the liver or bile system.
antigens. It arises mainly from the inactivation or death of binary specification A decision between only two alternative states.
immature potentially self-reactive B cells when they encounter binding constant Equilibrium constant for an interaction written
their specific antigen. as an association reaction.
Bchla Bacteriochlorophyll a. binding function Concentration of ligand bound divided by the
Bence-Jones protein Light chains of immunoglobulins produced total acceptor concentration.
in excess by a malignant monoclonal B-cell line (e.g. in multiple binding isotherm Mathematical expression of the relationship
myeloma). between ligand concentration bound to a macromolecule and the
benign paroxysmal positional vertigo Common form of vertigo free form of the ligand at equilibrium.
due to canalolithiasis (free-floating heavy debris from otoconia) binding pocket Putative region of a receptor where the ligand is
of the posterior semicircular canal. recognized and bound.
benthic Living primarily on the sea or lake floor, at the bottom of binding site Site on a macromolecule at which another molecule
the water column. binds specifically.
benzene ring The ring system composed of six carbon atoms. binomial nomenclature The system of naming organisms created
by Swedish naturalist Carolus Linnaeus, in which two Latin
benzynes A group of intermediates, unstable under most condi-
names are used, e.g. Homo sapiens. The first name is that of the
tions, that result from the removal of two hydrogen atoms from
genus, and the second is the specific name, which distinguishes
an aromatic hydrocarbon.
that species from other species in the genus.
Berlese–Tullgren funnel See Tullgren funnel. biochemical threshold The level at which the mutated DNA load
berry Fleshy fruit that does not split open. in a cell type causes an apparent biochemical defect.
Bertrand lens A supplementary lens in the optical path of the bioconversion The conversion of biomass (organic matter, mostly
microscope, between the objective and eyepiece(s), that allows the from plants) into food, feed, biogas or biofertilizer through the
front focal plane of the condenser and back focal plane of the activity of living organisms, generally fungi.
objective to be viewed by the eyepiece(s). biodiversity See biological diversity.
bet-hedging Life-history traits that act to reduce variance in bioethics The study of ethical issues within biology and the other
fitness induced by environmental variation. life sciences, healthcare and medicine.
b-glucan A polysaccharide composed of b-D-glucopyranosyl biofacies A rock unit that contains an assemblage of fossils
units. The term is used primarily with those polysaccharides characteristic of a particular environment.
in which the b-D-glucopyranosyl units are linked (1-6) and/ biofilm Layers of bacteria, often forming individual consortia, that
or (1-3) and not with cellulose in which the units are linked attach to fluid interfaces or, more frequently solid surfaces,
(1-4). forming microbial communities.

153
Glossary

bioimmuration Fossilization of soft organisms as moulds in blastic development Formation of a conidium by the enlarge-
calcareous matter secreted by cemented bivalves or tube-building ment of a conidial initial that is distinct from the conidiogenous
polychaetes. cell and enlarges before a septa separates the conidium from the
biolistic transformation Introduction of DNA into cells by high- conidiogenous cell.
velocity bombardment with small metallic particles coated with blastoconidium A budding cell borne on a pseudomycelium. Also
DNA. The term ‘biolistics’ is short for biological ballistics. Also called a blastospore,
called microprojectile bombardment. blastomere identity In some organisms, the descendants of a
biological community A group of one or more populations of given blastomere show a complex invariant differentiation
plants, animals and other organisms, living in a common space pattern with many different cell fates characteristic of that
(region). blastomere.
biological diversity The complete range of species, genetic blastomeres Cells formed by the first few embryonic cleavages.
variation, biological communities, and ecosystem processes on blastopore In sea urchins and amphibians, the site of movement of
Earth, or in a particular designated area. cells from the outside to the inside of the embryo during
biological species concept The idea that species are groups gastrulation.
of interbreeding natural populations that are reproductively blastula The hollow sphere of cells that results from the cleavage
isolated from other such groups. divisions in many animal embryos. It precedes the formation of
bioluminescence Production of light by living organisms. germ layers.
biomimetic matrix An artificial material able to duplicate the blastulation The development of a blastula.
biological properties of normal tissue extracellular matrix. bleaching Loss of symbiotic algae, and hence pigmentation, in
bioreactor A device designed to support the growth of mamma- corals and other symbiotic reef organisms in response to stress.
lian, plant, or microbial cells. Such a device typically provides
blepharoplast Basal body of flagellates, typically associated with a
contact between cells, growth medium and oxygen/air. It may
flagellum and sometimes additional structures. it is identical to
support the growth of either attached cells or cells in suspension.
the basal body or kinetosome.
bioremediation The use of organisms or biological products to
blood–brain barrier A selective mechanism that hinders the
decontaminate polluted ecosystems.
passage of cells and large molecules, including many antimicro-
biosurfactant An amphiphilic molecule of biological origin. Such
bial agents, from the bloodstream to the cerebrospinal fluid.
a molecule has the surface-active properties of being able to
Bloom syndrome (BS) Disorder characterized by growth retarda-
reduce surface and interfacial tensions. Microorganisms very
tion, facial erythema and proneness to respiratory tract
often make them to emulsify and solubilize n-alkanes.
infections.
biosynthesis The formation of a natural product in a living system.
blot A nitrocellulose or nylon sheet that contains spots of
biotic Of or pertaining to living organisms.
immobilized macromolecules (protein, DNA or RNA) or their
biotic factors The effects of living organisms on other living things
fragments. A molecular probe is applied to identify specific
in ecosystems.
components of the spots.
biotin–streptavidin A detection system that utilizes the high
affinity of streptavidin for biotin. For example, the binding of blotting General term for the transfer of protein, RNA or DNA
enzyme-conjugated streptavidin to biotinylated antibodies con- molecules from a relatively thick acrylamide or agarose gel to a
stitutes a convenient detection system in ELISA. thin membrane (usually nylon or nitrocellulose) by capillarity or
biozone A stratigraphic unit characterized by a fossil species or an electric field.
association. blue-light photoreceptor A protein that senses a blue-light signal
bipedal Describes animals for which a gait using the two hindlimbs and transduces this signal to downstream proteins that elicit the
only is a significant part of their repertoire of movement. biological responses to blue light.
bipedalism The ability to walk on the hindlimbs only. Many extant body fossil A fossil that preserves part, or all, of the body of an
primates can walk for short periods on their hindlimbs, but only ancient animal or plant.
modern humans are well adapted to do so. bolide A general term used for an extraterrestrial object that strikes
biramous limbs Arthropod limbs having two rami or branches, i.e. the Earth and explodes, without specifying if the object is a comet,
both endopod and exopod. meteor or asteroid.
bivalent A pair of synapsed homologous chromosomes, each bone marrow Highly vascular cellular substance in the central
having two sister chromatids, that appear during prophase and portion of many bones that synthesizes all the blood cells
metaphase I of meiosis. (erythrocytes, leukocytes and platelets).
bivalves A class of molluscs (phylum Mollusca, class Bivalvia (or bone marrow transplantation The correction of a lack (as in
Pelecypoda)) including clams, scallops and oysters, which have immunodeficiencies) or abnormality (as in leukaemia) of blood
bivalved, hinged shells, typically with left and right valves cells by extracting bone marrow cells from a healthy donor
showing mirror image symmetry, present from the Cambrian to and transfusing them into the patient to replace the defective
the present. bone marrow. Unless the patient is immunodeficient, the
BLAST Basic Local Alignment Search Tool, a sequence-based bone marrow must be matched for tissue type to avoid being
search tool developed at the National Center for Biotechnologi- rejected.
cal Information, that compares a given sequence with a database bone-lining cells The primary cell in the periosteal and endosteal
of sequences and produces an optimal local alignment. surfaces of adult bone, which are, in essence, inactive osteoblasts.
blastema Mass of proliferating mesenchymal cells that forms at book gills Gills of horseshoe crabs and some extinct chelicerates
the site of an amputated limb in some animals and can regenerate formed by a series of closely spaced cuticular plates (lamellae) and
the severed limb. located on appendages of the opisthosoma.

154
Glossary

book lungs Respiratory organs of certain terrestrial chelicerates buffer capacity A measure of the degree to which a buffer can resist
(scorpions, spiders, whipscorpions) formed by a series of closely a pH change. ‘Practical buffer capacity’ is the concentration of H1
spaced cuticular plates (lamellae). The lungs are contained within that must be added or removed to change the pH by one unit.
an air-filled chamber that opens to the outside through a hole. bulges/bulged-out nucleotides Nucleotides within a stem re-
Boolean operators When searching a database with two or more gion that are not base paired, i.e. are located between two helical
terms, the terms are combined by the operators AND, OR, NOT. segments.
The default Boolean operator is AND. bulla The bony covering of the middle ear space in mammals.
borders Well-defined sequences at the extremities of T-DNAs that bunodont The condition where cusps of the molar teeth are
are cut by the T-DNA processing enzymes. bulbous, as in humans.
boutons Small synaptic contacts on hair cells. buoyancy The upward force of any fluid upon an object immersed
BPA Brooks’ parsimony analysis, a technique for comparing host in it.
and parasite cladograms. byssus Solid mass or threads secreted by a bivalve mollusc to attach
BPI See bactericidal permeability-increasing protein. itself to a substrate. Sometimes the byssus is calcified, and in most
brachials The generally smaller, dorsal valve of a brachiopod shell, cases it can be renewed if destroyed.
which supports a lophophore (a tentacle-bearing feeding organ).
brachiation Swinging along branches by the arms.
bradycardia Severe slowing of the heart rate. C period The time for a round of DNA replication, from initiation
brain death A widely accepted standard for the declaration of at the origin to completion at the terminus. In Escherichia coli the
death based on the death of the whole brain. C period, for cells growing between 20 and 60 min doubling times,
brain plasticity Change in the output of the brain that depends on is a constant of approximately 40 min.
experience, and reflects changes in synaptic connections. C3 convertase Bimolecular proteolytic enzyme from the classical
brainstem That part of the brain located above the spinal cord and and alternative complement pathways. It cleaves complement
comprising the medulla, pons and midbrain. protein C3 into fragments C3a and C3b.
branch migration In general genetic recombination, a step during C3 pathway Photosynthetic pathway of carbon dioxide reduction
homologous DNA pairing and strand exchange in which the and incorporation into carbohydrate in which the first product is
length of the initial DNA joint is extended. the three-carbon compound phosphoglycerate. Most plants use
branch site The location within an intron at which a 20 ,50 the C3 pathway.
phosphodiester bond is formed during the first step of C3 photosynthetic carbon reduction cycle See C3 pathway.
splicing. C4 pathway Photosynthetic pathway in certain types of plant (C4
brassinosteroids Plant steroid hormones that are involved in plants) in which carbon dioxide is preconcentrated in a process
stem elongation and photomorphogenesis. An example is involving four-carbon compounds such as malate and aspartate
brassinolide. before fixation by the C3 reactions.
bronchoalveolar lavage fluid Fluid recovered from washings of C5 pathway A pathway for the synthesis of 5-aminolaevulinic
the lower respiratory tract. acid, a common precursor for tetrapyrroles, from glutamic acid
bronchodilators Drugs that relax airway smooth muscle, usually using tRNAGlu.
resulting in enlargement of the airway lumen, and improved C24 An ecotype of Arabidopsis thaliana.
bronchial air flow. Ca21-ATPase A transmembrane transport protein that hydrolyses
bronchospasm Constriction of airway smooth muscles, resulting ATP and uses the energy released to transport Ca2 1 across a
in narrowed airway lumen size, which reduces the amount of air membrane.
that can flow through the airways. cachexia General weight loss and wasting occurring in the course
browser Computer program used for navigation and for viewing of a chronic disease.
information on the World Wide Web. The interface is based on cadastral gene Any member of a class of floral homeotic genes that
highlighted hypertext links, active words, phrases and graphics repress the activity of other floral homeotic genes. They do not
that can be clicked on to jump to different information. have a positive function in specifying organ identity on their own.
brush border Microscopic projections of the luminal membrane of cadherins Family of proteins that mediate calcium-dependent
intestinal cells. cell–cell interactions in animal tissues.
bryozoan ‘Moss animals’ (phylum Bryozoa), marine and fresh- caecilians A group (order Gymnophiona) of living amphibians
water colonial organisms whose colonies are made up of characterized by very elongated limbless bodies and large
numerous tiny animals, each of which normally bears a copulatory organs in the tail region. They are tailless or nearly so.
lophophore, a crown of ciliated tentacles for filter feeding. caecum Blind-ended pouch in ruminants formed at the junction
Present from the Ordovician to the present. between the small and large intestine (equivalent to the
bubble contrast agent A preparation of gas bubbles used to rudimentary human appendix), which has the specific function
increase ultrasound scatter, and therefore echogenicity in the of housing microbes and semidigested food in order to augment
body. assimilation of metabolites.
buccal Associated with the foregut (pharynx region). calcite A common mineral constituent of some animal skeletons
budding compartment The cellular location of virus assembly (e.g. brachiopods, bryozoans, many extinct corals, some clams).
and budding. It is composed of calcium carbonate (CaCO3), has a rhombo-
buffer Something that resists change. A pH buffer resists a change hedral or hexagonal crystal symmetry and is more stable than the
in the pH of its solution when small quantities of H1 are added (or other polymorph of calcium carbonate, aragonite. It is the main
produced therein) or consumed. constituent of limestone.

155
Glossary

calcium inhibition Inhibitory effect of Ca21 on actin–myosin carbenes A group of chemical intermediates, unstable in most
interaction, which causes cytoplasmic streaming of plasmodia instances, that contain a divalent carbon atom.
and amoeboid movement of amoebae. This action of Ca21 is carbocations A group of chemical intermediates, unstable under
ubiquitous in plants and lower eukaryotes but not in animals. most conditions, that result from the departure of an anion from
calix The Latin word meaning ‘cup’ from which the name an organic molecule, or from its protonation.
calicivirus was derived. It refers to cup-like structures on the carbon cycle The cycle of carbon on Earth through atmosphere,
surface of the virus particle that are characteristic of calicivirus living organisms, oceans and rock.
morphology. Carboniferous period The penultimate major division of the
callase An enzyme that degrades the wall of the pollen tetrad and Palaeozoic era.
releases the pollen grains. carboxylation In plants that photosynthesize via the C4 pathway,
callus Unorganized mass of cells that forms on the surface of a the addition of a carbon dioxide molecule to phosphoenol-
wound in plants, or when a piece of plant tissue is cultured. pyruvate to form four-carbon acids in mesophyll cells.
calnexin Protein found in the endoplasmic reticulum that acts as a carboxysomes Primitive carbon dioxide-fixing organelles present
molecular chaperone for glycoproteins and assists in protein- in some bacteria.
folding pathways. carcinogenesis The generation of a cancer. The process by which a
calorimetry Measurement of the amount of energy (heat) liberated normal cell is converted into a cancer (tumour) cell.
by metabolism. carcinization The evolution of a crab-like form.
calotte (1) The anterior or head region of dicyemid mesozoan carcinogen Any chemical or physical agent that can cause cancer.
parasites, by which they attach to the cephalopod kidney. (2) In Most carcinogens act by producing mutations in a cell’s DNA,
vertebrate anatomy, the brain case, that is the cranium minus the eventually leading to the transformation of this cell into a tumour
face and the cranial base. cell.
calreticulin Protein found in the endoplasmic reticulum that acts carcinoma A malignant tumour of epithelial cells. This is the most
as a molecular chaperone for glycoproteins and assists in protein- common type of cancer.
folding pathways. cardenolides Derivatives of the plant steroid calotropagenin,
calvaria In anatomy, the cranium minus the face. which has the lactone ring at position 17, a characteristic of
cardiac poisons.
calyx (1) The outermost part of a flower, consisting of the sepals.
cardiac arrhythmias A group of cardiac disorders characterized
(2) Contacts of sensory neurons that wrap around the entire
by irregular cardiac rhythm. There are two types of arrhythmias:
hair cells.
tachycardia (the heart beats too rapidly) and bradycardia (the
CAM See crassulacean acid metabolism.
heart rate is too slow).
cancer A general term for any malignant growth. The word derives
cardiac pacemaker A region of the heart where specialized cardiac
from the Latin for ‘crab’, as many tumours resemble a crab, with
cells spontaneously depolarize and set the rhythm and frequency
the body represented by the main tumour mass and the claws by
of the heartbeat.
the invasive tumour margins.
carinate Applied to birds having a keel on the sternum (breastbone).
candidiasis Invasive infection caused by species of the fungus carnivore An animal that eats other animals.
Candida. carotenoids Orange, red and yellow terpenoid pigments in living
canonical structure A small repertoire of main-chain conforma- organisms, that have a photoprotective and structural role.
tions observed in the antigen-binding sites of antibodies. Most carpel The female reproductive structure in angiosperm flowers
known variations in the amino-acid sequences of antigen-binding that encloses the ovule(s). Carpels can occur either separately or
sites modify the surface by altering side-chains on these main- fused with other carpels.
chain structures. carrageenan A sulfated galactose and anhydrogalactose-rich
cap A methyl guanosine found at the 50 end of some RNAs and polysaccharide in the walls of red algae, especially the order
joined to it by an unusual 50 ,50 triphosphate linkage. All RNAs Gigartinales. It is extracted and used as a gelling agent and
transcribed by RNA polymerase II (including pre-mRNA), most stabilizer in the food and pharmaceutical industries.
spliceosomal snRNAs and several other snRNAs are capped. cartilage Tough but flexible skeletal tissue composed of an organic
capping (1) The formation of a methylated GTP on the 50 end of matrix of chondrin.
eukaryotic mRNA and some other RNAs during their matura- caruncle A fleshy protuberance.
tion. (2) Binding of a protein to the end of a microfilament of cascade Organization of multiple enyzmes in succession within a
microtubule. signalling pathway, each activating the next enzyme in the pathway.
capsid The protein coat surrounding the nucleic acid core of a case–control study A study in which it is determined whether
virus. The proteins are virus encoded. an exposure to some factor (e.g. increased plasma cholesterol
capsid protein A protein component of the coat or outer shell of concentration) is different between cases (e.g. individuals with
the virus particle. myocardial infarction) compared with controls (individuals
capsule (1) Multiple-seeded fruit of a syncarpous ovary that breaks without disease).
open by splits or pores. (2) Highly hydrated amorphous material, case-fatality The proportion of all cases of a disease who die.
chiefly polysaccharide, which forms a thick outer coat, outside Casparian band Modifications (containing suberin and lignin) in
the cell wall, on some bacteria. the radial cell walls of the root endodermis.
captive breeding The breeding of plants or animals in captivity, caspases A family of aspartate-specific cysteine proteases that
usually with the aim of reintroducing populations into the wild. mediate the execution phase of cell apoptosis. Caspases are
carapace Outgrowth of head ectodermal tissue in arthropods, related to mammalian interleukin 1b-converting enzyme (ICE/
covering and often fusing to the thoracic segments. caspase-1) and to the nematode apoptotic gene product Ced-3.

156
Glossary

catabolic Degradative. CD34 A cell-surface protein present on a small population of


catabolism Metabolic reactions in which substrates are broken haematopoietic cells that includes stem cells and multipotential
down, liberating energy. progenitors.
catalase Enzyme that removes potentially toxic hydrogen peroxide CD4 Cell-surface protein that distinguishes a major subset of T
(H2O2) from cells by degrading it to water and molecular oxygen. cells. It acts as a co-receptor in conjunction with the T-cell antigen
In animal cells, catalase is found mainly in peroxisomes, and is receptor by binding to MHC class II molecules.
frequently used as a marker for the peroxisomal compartment. CD4 T lymphocytes, CD4 T cells One of the two major subsets of
The ‘catalase test’ is used widely in microbiology as a diagnostic T lymphocytes, distinguished by the presence of the co-receptor
test. molecule CD4 on the cell surface. CD4 T cells include cells that
catalysis Increase in the rate of a chemical reaction achieved activate infected macrophages and the ‘helper’ T cells that
by a substance, the catalyst, that remains unchanged after provide antigen-specific ‘help’ to activate B cells and other T cells.
the reaction. In catalysis, the rate of reaction is increased CD46 Decay-accelerating factor, a cell-surface protein that pro-
without altering the position of the overall chemical tects cells from lysis by complement, and also serves as the
equilibrium. receptor for measles virus.
catalysis of elementary steps An increase in reaction rate that CD8 Cell-surface protein that distinguishes a major subset of T
results from an interaction of the enzyme and substrate that cells. It acts as a co-receptor in conjunction with the T-cell antigen
selectively stabilizes the transition state of the chemical catalytic receptor by binding to MHC class I molecules.
step. CD8 T lymphocytes, CD8 T cells One of the two major subsets of
catalyst Any substance that speeds up a chemical reaction while T lymphocytes, distinguished by the presence of CD8 on the cell
remaining unchanged at the end of the reaction. Enzymes are surface. CD8 T cells are mostly cytotoxic cells responsible for
catalysts. the antigen-specific killing of infected or foreign cells; they may
catalytic power The effectiveness of a catalyst, as measured by the also be involved in inhibiting the activity of other T cells. Also
rate enhancement that it causes. called cytolytic T lymphocytes (CTLs), or cytotoxic or killer
catalytic RNAs RNAs that are able to catalyse, in the absence of T cells.
proteins, specific chemical reactions. For natural RNAs, these CD26P See P-selectin.
reactions are usually transesterifications affecting the catalytic CDK See cyclin-dependent kinases.
RNA itself. cDNA A DNA of sequence complementary to an RNA. It can refer
catalytic triad A constellation of functional groups at the active either to a single-stranded DNA copy of the RNA, or to the
site of an enzyme that can work in cooperation to make a reaction double-stranded form of this DNA. Synthesis of cDNA on the
proceed more quickly than is possible with any single function RNA template is catalysed by reverse transcriptase.
acting alone. cell banking Preservation of cells in a frozen state.
catarrhine A general term of primate classification that refers to cell body The part of a neuron containing the nucleus, ribosomes
the Old World monkeys, apes and humans. and other cytoplasmic components required for cell survival and
catecholamine A compound consisting of a catechol nucleus (an maintenance.
aromatic structure with two adjacent hydroxyl groups on the cell cycle stages The intervals between one mitosis (M) and the
benzene ring) and hydrocarbon tail ending in an amine group. next are known as gap 1 (G1), DNA synthesis stage (S) and gap 2
The body’s endogenous catecholamines are noradrenaline, (G2).
adrenaline and dopamine. cell cycle The sequence of stages that an actively growing cell
category A level such as the genus in a formal classification system. passes through between the time it is formed and the time it
catenane Interlinked double-stranded DNA circles, as in the links divides to give two new cells. During this time it doubles its
in a chain. cytoplasmic constituents, replicates its DNA, and finally divides
caudal Towards the tail. to give two daughter cells. The stages are distinguished by the
caveolae Bulb-shaped invaginations at the plasma membrane state of the nuclear DNA within the cell. The cell cycle’s four
of ~50 to ~100 nm diameter, distinguished by the presence of phases are G1 (first stage of interphase), S phase (period of DNA
oligomers of the protein caveolin and enriched in cholesterol and synthesis), G2 (postsynthetic interphase) and M (mitosis). Also
sphingolipids. They are involved in the transport of small called the cell division cycle.
molecules into the cell. cell envelope All structures outside the cytoplasm in bacteria. It
cavitary lesions An abnormality on a chest radiograph showing a consists of the inner (cell) membrane, the peptidoglycan cell wall,
localized infiltrate with a central lucency (hole) in an otherwise the outer membrane (in Gram-negative bacteria), and the
dense infiltrate. capsule, when present.
CD marker See CD nomenclature. cell extract The soluble constituents of cells, obtained by breaking
CD molecule See CD nomenclature. cells open and separating out the insoluble material.
CD nomenclature The CD (cluster differentiation) nomenclature cell fate specification The point in a cell lineage at which a cell
system is used to identify specific antigenic structures on blood becomes fixed, or determined, in the type or types of cells and
leukocytes and tissue cells. A cell-surface protein that can be tissues its descendants can produce.
identified uniquely by a set of monoclonal antibodies is cell homogenate Buffered suspension containing all cell orga-
designated CD followed by a number, e.g. CD4. CD molecules nelles and subcellular components after the mechanical destruc-
are often used as markers to distinguish different cell types and tion of tissues or cells.
stages in differentiation. cell invasion Penetration of cells through basement membranes
CD3 A component of the T-cell receptor complex. and extracellular matrix.

157
Glossary

cell junctions Structures that hold the faces of neighbouring cells Cenozoic Geological era from 65 million years ago to the present.
together, especially in an epithelium. Epithelial cell junctions central cell Cell in the embryo sac of an angiosperm that fuses with
include tight junctions and desmosomes. Other types of cell one sperm to give rise to the endosperm, a nutritive tissue for the
junction are gap junctions, by which cells can communicate with developing embryo.
each other, and septate junctions in insect cells. central dogma The information for the amino-acid sequence of a
cell lineage A pedigree of cells produced from an ancestral cell by protein resides in the nucleotide sequence of the RNA, which is in
mitotic divisions. Cells may adopt specific developmental fates turn encoded by the nucleotide sequence of the genes in DNA; in
because of their unique lineages. short, ‘DNA makes RNA makes protein’.
cell membrane In bacteria, usually refers to the lipid bilayer central nervous system (CNS) The brain and spinal cord.
membrane immediately surrounding the cytoplasm and under central pattern generator An oscillatory neural network that
the peptidoglycan cell wall. It is analogous with the plasma produces a stereotypical rhythmic motor output.
membrane of eukaryotic cells. Some types of bacteria have an centric fission Reverse process of centric fusion.
additional outer bilayer membrane containing lipopolysacchar- centric fusion A genetic process whereby two acrocentric
ide, outside the peptidoglycan layer. chromosomes (in which the centromere is located near the
cell migration Movement of cells over a surface. end of the chromosome) are joined in the region of their
cell models Cells that have been permeabilized by mild detergents centromere.
and subsequently incubated in experimental solutions, usually in centrifugal Away from the axis or centre.
order to reactivate a living process, e.g. motility. centring telescope A telescopic eyepiece that can be inserted in
cell signalling Communication between components of an in- place of an eyepiece to permit alignment of the phase-contrast
dividual cell or between cells through the generation and release microscope. An alternative to the Bertrand lens.
of bioactive compounds. centriole A specialized, self-replicating microtubule-organizing
cell transformation A change to a state of unrestricted growth in centre. In animal cells, the mitotic spindle microtubules form
culture, resembling the tumorigenic condition. between a pair of centrioles.
cell wall In plants, algae, fungi, bacteria, and archaea, the semi- centripetal Towards the axis or centre.
rigid layer of material lying immediately above the plasma centromere Specialized constricted region of a chromosome at
membrane. In plants and some protists, the wall is composed which the pair of chromatids in a metaphase chromosome are
mainly of cellulose. Fungal cell walls are composed mainly of held together. The centromere divides the chromosome into two
chitin. In bacteria, the cell wall consists chiefly of peptidoglycan. arms. It contains the kinetochores, which attach the chromosome
Walls of Gram-positive bacteria consist of a thick layer of to the spindle and ensure correct segregation of chromatids
peptidoglycan plus secondary polymers such as teichoic acid. during mitosis and meiosis.
Gram-negative walls consist of a thin layer of peptidoglycan, over cephalic version The turning of the baby in utero by external
which is an outer lipid bilayer membrane containing lipopoly- manipulation so as to change the position (usually from breech) in
saccharide. In archaea, the cell wall can be a simple S-layer, a order to have the head of the baby as the presenting part for delivery.
multi-S-layered complex, or an amorphous matrix of pseudo- cephalon The head of a trilobite, often semicircular in form, and
murein or methanochondroitin. usually with compound eyes.
cell-mediated cytotoxicity The direct killing of one cell by ceramide A basic building block of all the complex glycosphingo-
another. lipids. It consists of a sphingosine base in which the amino group
cell-mediated immunity Immunity mediated by the T lympho- is N-acylated with a long-chain fatty acid.
cytes of the immune system. It can be transferred to naive cerci Abdominal appendages of insects.
individuals by transferring cells from an immunized individual cerebellum An upward hemispherical outgrowth from the hind-
but not by plasma or serum. brain which plays an essential role in motor coordination.
cell–substratum contacts Sites on a cultured cell at which it is cerebral cortex Outer layer of grey matter covering the cerebral
attached to a solid substratum. These sites are distinguished by hemispheres.
the formation of specialized cytoskeletal structures. cerebrum The most anterior part of the brain, also called the
cellular immunity See cell-mediated immunity. telencephalon.
cellular oncogene In a eukaryotic cell, a gene that, if activated by a cerophyl A form of dry, highly refined, powdered grass
virus or mutation, can cause the cell to become a tumour cell and leaves available commercially. It stimulates the growth of
develop into a tumour. bacteria, which can then be used as food by ciliates or other
cellular P450 system Membrane-bound enzymes present in the protists.
endoplasmic reticulum that convert nonpolar chemicals to water- CFTR Cystic fibrosis transmembrane protein.
soluble, excretable forms. Chagas disease Disease caused by Trypanosoma cruzi, restricted
cellular responses See cell-mediate immunity. to the New World.
cellulolytic Able to digest cellulose molecules into simpler carbo- chancre The primary infectious syphilis lesion which occurs at
hydrates. the site of inoculation of infection in a person with no
cellulose A linear polysaccharide polymer of b-D-glucose, which is previous infection. It is always associated with dependent
deposited in the form of crystalline microfibrils and forms the lymphadenopathy.
main component of plant cell walls. channels Transmembrane proteins that function as selective pores
cement gland An ectodermal derivative found in amphibian in the membrane to allow the passage of ions and other solutes.
embryos just anterior to the neural plate. It arises as a result of Channels are usually gated, opening as a result of a particular
neural induction. The gland secretes a sticky mucous substance stimulus to let an unlimited number of ions or other solutes
that allows embryos to anchor themselves to aquatic plants. through, and then closing again.

158
Glossary

chaperone A protein that interacts with unfolded, often newly chemotherapy-induced neutropenia A deficiency of blood
synthesized proteins, and assists in their folding to the native neutrophils due to cancer chemotherapy drugs that suppress
state. Chaperones promote folding mainly by suppressing the production of neutrophils in the bone marrow.
aggregation of unfolded proteins with one another. Chaperone chert Rock made of extremely fine-grained quartz silicon dioxide.
proteins include GroE, Hsp70 and Hsp90. They are essential for Different colour varieties are known as flint or jasper.
life and are abundant in all cells. chi Crossover hotspot instigator, the DNA recognition sequence
chaperonin Any of a class of chaperone proteins that participate in (50 GCTGGTGG30 ) for the bacterial RecBCD enzyme. Chi
protein folding. modulates RecBCD activity into a recombination mode, and
chaperone/usher pathway A process in the assembly of bacterial from double-strand exonuclease mode, which results in elevated
fimbriae that requires a periplasmic protein (the chaperone) that recombination at and near Chi sites.
helps fold pilin protein molecules and an outer-membrane protein chimaera (1) Embryo derived from a composite of two or more
(the usher) that directs the ordered assembly of pilins. zygotes. (2) Any organism that is a composite of cells of two or
characterization The process of obtaining data on organisms. more genotypes, for example, experimental mouse bone marrow
charging See aminoacylation. chimaeras are produced by killing the bone marrow and replacing
Chédiak–Higashi syndrome (CHS) A disease characterized by it with bone marrow of a different genotype.
decreased pigmentation, abnormal susceptibility to pyogenic chimaeras Cartilaginous fishes related to sharks and rays.
infections and neurologic disorders. chimaeric (1) Describes any organism that is composed of cells of
checkpoint A point in the cell cycle where progression through the two or more different genotypes. (2) Describes a DNA molecule
cycle is stopped while errors are corrected. containing DNA from two or more different sources, e.g. a
chelation The formation of a coordination compound between a recombinant DNA.
metal cation and an anionic ligand. chimaeric chromosomes Chromosomes made by joining
chelicera (plural chelicerae) The first appendage in chelicerates together DNA from two or more different chromosomes.
(spiders and their allies). It is typically composed of two or three chimaeric mice Mice composed of cells of two different genotypes.
segments and is specialized for capturing and manipulating food. Such mice may be produced by transgenesis or by the fusion of
cheliped In arthropods, a walking leg equipped with a claw. very early embryos from two different sources.
chemical potential (ll) The rate of change of the Gibbs energy (G) chimaerism The situation in which cells from genetically different
of a system with respect to the amount of species i at constant individuals coexist in one body.
temperature (T), pressure (P) and the amounts of other species chirality The ‘handedness’ of a molecule, so that it is not super-
present. It is used in derivations, but is replaced by DfGI in imposable on its mirror-image.
calculations. chitin A high relative molecular mass polymer of 1:4 linked N-
chemoattractant A chemical that attracts bacteria (or other cells acetyl glucosamine residues.
or organisms) to move towards it. chitinous Containing the polysaccharide chitin (poly-N-acetyl-
chemocline The zone between two stable and long-term water glucosamine).
layers of different chemistry and specific gravity. chitinozoans Extinct tiny flask or bottle-shaped organisms
chemoheterotrophic growth Growth of an organism on organic believed to have been single-celled protistans, and which were
compounds, without the use of light as a source of energy. an important element of the Devonian zooplankton.
chemokines A superfamily of small secreted proteins that attract, chlamydospore A thick-walled asexual spore that develops from a
guide and activate white blood cells during immune reactions. transformed hyphal cell.
chemokinesis An increase in the random motion of a cell in chloride equilibrium Unequal distribution of chloride on the
response to a chemical stimulus. intracellular and extracellular sides of a cell membrane at which,
chemolithoautotroph Microorganism that obtains energy because of the presence of other nonpermeable ion species and the
through chemical oxidation, using inorganic compounds as resting potential, no chloride flux occurs.
electron donors, and obtains its cellular carbon from the chloroplast Subcellular chlorophyll-containing organelle in higher
reduction of carbon dioxide. plants and algae in which light capture and the fundamental
chemolithoautotrophic Growing on inorganic carbon and energy transformation processes of photosynthesis occur. It
energy sources. contains a DNA genome which is transcribed in situ and encodes
chemo-organotrophic Using organic material as the source of some of the proteins essential for photosynthesis.
both energy and carbon for growth. chlorosis The yellowing of photosynthetic tissues associated with a
chemorepellant A stimulant that causes bacteria (or other cells or loss of chlorophyll.
organisms) to move away from it. chlorosome The nonmembranous light-harvesting structure in
chemosensation Sensory mechanism whereby an organism green bacteria.
recognizes chemicals (water or airborne) in its environment. choanocyte A cell type in sponges. It has a collar of cytoplasmic
chemotactic factors Molecules that induce chemotaxis or chemo- tentacles and an apical flagellum and is responsible for generating
kinesis. the water current through the sponge. Choanocytes are typically
chemotactic receptors A subfamily of seven-span G-protein- arranged in groups, making up a choanocyte chamber.
coupled receptors which bind chemokines and transmit the choanoderm The inner, or feeding, layer of the body wall of a
chemotactic signal to the cells. They are found mainly on sponge, composed of choanocytes.
leukocytes. cholera Acute gastrointestinal illness resulting from infection with
chemotaxis Orientation of an organism or cell towards the source the bacterium Vibrio cholerae. It is characterized by profuse,
of a chemical and its directional movement up a concentration watery diarrhoea and vomiting, resulting in severe, life-threaten-
gradient of the chemical. ing dehydration.

159
Glossary

cholestasis A stoppage in the flow of bile through the biliary tree, chromosome A thread-like structure that carries the genetic
before its excretion into the duodenum. information in eukaryotic cells. Chromosomes are contained in
cholinergic synapse A synapse at which acetylcholine (ACh) is the the nucleus, and each chromosome is composed of a single long
neurotransmitter. DNA molecule complexed with protein. Chromosomes only
chondrite A type of stony meteorite that contains numerous small become visible under the light microscope during mitosis and
spherules of silicate mineral. A subset, the carbonaceous meiosis, when they become highly compacted.
chondrites, contain several per cent organic carbon. chromosome elimination Elimination of whole chromosomes
chondroblast Specialized cell that secretes cartilage. Also known during mitosis, meiosis or interphase.
as a chondrocyte. chromosome inversion Type of chromosomal rearrangement in
chordate A member of the phylum Chordata, which includes which a segment of chromosome is inverted and the order of the
vertebrates and a few marine invertebrate groups such as genes on it reversed.
tunicates and amphioxus. chronic condition A long, continued illness.
chorion The proteinaceous outer shell of many arthropod eggs. chronic granulomatous disease (CGD) Immunodeficiency dis-
chorionic villus sampling Biopsy of the placenta. ease resulting from a defect in phagocytosis in neutrophils,
macrophages and other myeloid cells.
chromatid One of the two identical copies of a chromosome after
chronostratigraphy The geological time scale as defined by
replication, while the two copies are still held together at the
absolute time rather than the relative time scales of lithological
centromere.
and biostratigraphical units.
chromatin Originally defined by Flemming in 1880 as the deeply
chylomicron Droplet of fat present in the blood or lymph after
staining material present in the interphase nuclei of eukaryotic
absorption from the intestine.
cells and corresponding to the chromosomes. The term is now
chylous ascites Ascitic fluid which contains milk-white lymph
used more specifically to refer to the complex of DNA (and small
(i.e. chyle).
amounts of transcribed RNA) and proteins (histones and non-
chyme Pulpy acidic fluid consisting of gastric juices and partly
histone proteins) of which chromosomes are composed.
digested food that passes from the stomach into the intestine.
chromatin diminution Chromosomal fragmentation, followed by
cilia (singular cilium) Tiny motile hair-like structures present on a
the elimination of part of the chromosome during mitosis.
variety of eukaryotic cells, including some sensory receptor cells,
chromatin elimination Originally defined by Seiler in 1914 as and microorganisms. They are microtubule-based, containing a
the elimination of chromatin during the meiotic divisions. This core of microtubules in a stereotypical arrangement. In different
unfortunate term should be replaced by ribonucleoprotein cell types, cilia are involved in cell movement, sensory transduc-
shedding. The term is now used to denote the loss of chromatin tion, or the movement of fluid over the cell surface.
in chromatin diminution. circadian clock A self-sustaining oscillator system within an
chromatin marking system A system of enzymatic machinery in organism with a period of about 24 hours, and occurring
the cell that blocks expression of certain segments of DNA by independently of external synchronizing signals. It organizes
attaching additional chemical groups to some base pairs in various cellular and physiological processes with the day–night
those segments. Chromatin marks are transmitted in cell cycle.
lineages to create differential gene expression patterns in different circadian rhythm Any rhythmic biological process within an
cells. organism with a period of about 24 hours and occurring
chromatography The separation of molecules of different sorts by independently of external synchronizing signals.
dissolving them in an appropriate solvent and passing them over a Circe A sorceress who lured Ulysses and his men into her palace
solid support of various types (e.g. paper or a column of packed and turned them into pigs.
resin beads). Molecules will be differentially retained on the circular permutation The variable location of DNA termini in a
support on the basis of their size, charge or binding properties, virus population.
depending on the type of chromatography used. circularization End-to-end joining of linear DNA molecules to
chromophore A chemical group that can absorb light and form a circular DNA.
undergo a conformational change. Such groups mediate bio- circularizing oligonucleotide probes Oligonucleotides that
logical responses to light. A chemical group that becomes light- when successfully hybridized can be converted to closed
absorbing in the course of a chemical or enzymatic reaction is molecules ligated to the target sequences. Also known as ‘padlock
called a chromogenic group. probes’.
chromosomal polymorphism The presence of more than one circularly permuted DNA A collection of DNA molecules whose
type of chromosomal morphology in a population. sequences could all be derived from the same circular DNA
chromosomal rearrangement General term for any chromoso- molecule by cutting it (once) at different points.
mal abnormality in which part of the chromosome has been circulative transmission Persistent transmission of a pathogen
deleted, duplicated, inverted, transposed to another position on during which the pathogen does not replicate in the vector, but is
the chromosome, or translocated to another chromosome. passed through both the digestive and circulatory systems of the
chromosomal speciation Speciation as a result of chromo- vector.
somal rearrangements that cause post-mating reproductive circumventricular organs Epithelial organs that line the ventricle
isolation. of the central nervous system. They are typically composed of
chromosomal translocation Relocation of a chromosomal seg- neuronal and nonneuronal elements.
ment within the chromosomal complement of any cell, usually cirri (singular cirrus) Clusters of individual cilia acting as a unit in
involving a reciprocal exchange of material between two or more locomotion.
chromosomes. cis On or affecting the same molecule.

160
Glossary

cis-acting Describes a gene or regulatory element that affects genes clonal rearrangement The case where a large number of
located on the same chromosome. lymphocytes contain an identical rearrangement of the
cis-acting determinant Describes a component of a DNA/RNA immunoglobulin or T-cell receptor genes, usually as a result of
sequence that influences the activity or level of the same proliferation of the cell containing the original rearrangement.
sequence. For example, the 30 stem–loop structure that clonal selection theory The theory underpinning our present
protects an mRNA against exoribonucleases is a cis-acting understanding of the adaptive immune response. It requires that
determinant. individual lymphocytes each have a single unique antigen
cis-acting element A site in the regulatory region of a gene that specificity, and that only those lymphocytes whose receptors
usually delineates a site of interaction with a gene-regulatory match epitopes on the antigen will be activated to proliferate and
protein. achieve effector status. Originally formulated as a theory of
cisternae (singular cisterna) Membrane-bounded disc-shaped antibody formation by F. Macfarlane Burnet.
sacs that stack to form each individual Golgi body or cisternal clonal selection The activation and proliferation of small pre-
stack. existing clones of antigen-specific lymphocytes during the course
cistron A DNA sequence encoding a single polypeptide chain or a of an adaptive immune response.
single functional RNA molecule. clonal tumour Malignancy in which all cells share a single
clade A group of species that derives from a single ancestor common genetic ancestor.
(monophyletic) and includes all descendants of that ancestor. clonality The evolution and spread of a single gene, a single group
clade rank In a clade of N taxa, each taxon has an age rank from 1 of genes or a single strain of a bacterium locally or worldwide in
to N, reflecting the order of first appearances (with the oldest response to environmental changes.
given the rank of 1 and the youngest given a rank of N ). clone (1) A group of cells that are all descended from the same
cladistic Phylogenetic ordering of organisms based on the evalua- progenitor cell and thus have the same genetic make-up. (2) A
tion of individual characters. group of individuals, derived by asexual reproduction, that are
cladistics A method of working out the phylogeny of organisms by genetically identical to the parent and each other. (3) A fragment
building phylogenetic trees, or cladograms, that group organisms of DNA isolated from the genome and produced in multiple
strictly on the relative recency of common ancestry. copies by introducing it, via a specific vector, into a living cell
cladogenesis The generation of new lineages (clades) by specia- where it will reproduce itself. (4) To isolate and replicate an
tion. individual gene.
cladogram In phylogenetic systematics (cladistics), a tree-like clone contig A set of overlapping DNA clones.
representation of a phylogenetic hypothesis based on shared closed circular DNA Double-helical DNA in which both strands
apomorphies (synapomorphies) of the taxa included. It depicts are closed circles; i.e. there are no free ends.
basic (i.e. sister taxon) relationships among taxa but lacks cluster headache A disorder, mainly of men, characterized by
information about more specific (i.e. ancestor–descendant) agonizing, daily, one-sided, intense pain lasting for 30–120 min,
relationships. accompanied by a running eye, blocked nostril and intense
Clark electrode Electrochemical cell used for the determination of restlessness. The cluster of daily attacks ceases after 4–12 weeks.
oxygen concentration. Cluster of differentiation (CD) markers See CD nomenclature.
classification Ordering of organisms based on a given theory and cmc See critical micelle concentration.
philosophy. C–N glycosidic bond Covalent link between C1 of the ribose ring
class switch See isotype switch. and N3 of pyrimidine or N9 of purine heterocyclic rings.
clastogen A chemical capable of breaking chromosomes. cnidocyst A stinging organelle in cnidaria. It is a capsule-bearing
clathrin-coated pits Regions of plasma membrane that are inverted tubule, sometimes capped by an operculum. Pressure on
susceptible to sequestration, or endocytosis, due to cytoplas- a fine-hair-like cnidocil triggers its eversion, which in some cases
mic-side association with clathrin. is accompanied by the release of toxins.
claudication Occlusion of blood vessel flow, usually caused by CNS stimulant A substance that stimulates activity in the central
atheromatous plaque formation, leading to ischaemia of tissues. nervous system (i.e. the spinal cord and the brain).
cleidoic egg An enclosed egg incorporating much yolk to nourish CNS See central nervous system
the developing embryo. Also called an amniotic egg. coagonist Chemical that is required for receptor activation in
climbing fibre Axon of an inferior olivary neuron forming addition to the principal neurotransmitter.
multiple synaptic junctions with a Purkinje cell; each Purkinje coagulation A defensive immunological and haemostatic process
cell receives one climbing fibre. in which the blood solidifies locally, immobilizing the foreign
cline Variation in a trait along an environmental gradient or across antigen and preventing excessive bleeding following injury. It
ecotones. involves a cascade of activity of protein coagulation factors. This
clionid A group of marine sponges (phylum Porifera) that results in the formation of an insoluble fibrin meshwork which,
perforate (bore into) skeletons of other organisms, making a together with blood platelets, forms a clot.
gallery of holes in which they reside. coagulocyte A synonym for granulocyte in arthropods.
cloaca In some animals, a common chamber connected to the coalescence The genealogy of a sample of homologous DNA
exterior that receives products from the digestive, urinary and sequences from a population. Looking forward from the
reproductive tracts. ancestral sequence they form a tree of descent, while looking
clonal expansion The multiplication of a single cell through backward from the presence they undergo coalescence events.
several divisions to produce a number of identical daughter coat protein Protein that encases the nucleic acid core of a virus
cells. particle.

161
Glossary

coatomer Complex of proteins forming a coat around COPI coleoptile A long, cylindrical, modified leaf acting as a sheath to
vesicles which are involved in retrograde transport across the protect the young shoot of a grass seedling.
Golgi stack and to the endoplasmic reticulum. Col The Columbia ecotype of Arabidopsis thaliana.
cobalamin Vitamin B12. collagen Strong, triple-helical fibrous protein that is the major
coccidioidomycosis Invasive infection caused by the fungus extracellular structural component of most connective tissues
Coccidioides immitis. including bone and cartilage.
coccolith A scale-like structure on the surface of prymnesiophy- collaterals Output connections made from a neuron that act in
cean golden algae. it consists of an organic plate surrounded by a parallel with, and secondary to, the main output connections.
rim of calcium carbonate crystals. collectins Family of proteins that contain a collagen-like segment
coccolithophores, coccolithophorids A group of microscopic attached to a lectin-like domain that binds carbohydrate. Several
photosynthetic planktonic algae that produce calcareous cocco- collectins play an important part in innate immunity.
liths on their surface. Coccolithophores first appear in the fossil collenchyma Peripheral supporting tissue in plants with thick-
record in the Late Triassic but they did not become an abundant walled cells.
component of the world’s plankton until the Cretaceous. colloid A stable mixed phase in which one substance (e.g. a solid) is
cocultivation The culturing together of a mixture of cells of nonhomogeneously distributed within another (e.g. a liquid).
different origins or from different individuals, generally for the colonocyte Cell of the epithelial lining of the gut wall.
purpose of transferring and propagating viral infection. colony-stimulating factor 1 (CSF-1) Also known as M-CSF, this
code for To specify the amino-acid sequence of a protein by the was the first haematopoietic growth factor to be purified and
sequence of nucleotides comprising the gene for that protein. named. It predominantly stimulates the growth of macrophage
codominant Describes different alleles that, when both are present (M) colonies.
at a locus, result in a phenotype different from that produced if columella An extension of the sporangiophore which occurs
only one type is present. within a sporangium.
codon A sequence of three consecutive bases in mRNA that combinatorial biosynthesis The biosynthesis of ‘unnatural’
encodes a particular amino acid. natural products by genetic engineering, e.g. by the introduction
coelenteron Digestive and distribution cavity of coelenterates. It of a gene for a particular pathway from one species into a different
acts as a hydrostatic skeleton and sometimes as a brood chamber species.
for planula larvae. combinatorial specification The idea that a combination of
signals or lack of signals specifies the fate of a cell.
coelom In certain animals, a fluid-filled cavity that lies between the
commensal An organism that benefits from its relationship with a
gut and outer muscle layers (the body wall) and is lined with
host organism but without either benefiting or harming the host.
mesodermally derived tissue.
commensalistic interaction Interaction in which one of the
coelomocyte The haemocyte (immunocyte) of annelids.
interacting species increases its fitness while the other species is
coenocyte A multinucleated cell. The plasmodium of slime moulds
not harmed or benefited as a result.
is a coenocyte.
comminution The conversion of leaf litter to organic topsoil by
coenzyme, cofactor A small organic molecule (often a vitamin) decomposers.
that assists the enzyme catalysis of certain classes of reaction (e.g.
commissural axons Axons that cross from one side of the body to
reduction of a ketone to an alcohol).
the other.
coevolutionary hotspots Local communities where an interac- commissure Axon bundle connecting the right and left halves of
tion between two or more species is under reciprocal selection (i.e. the brain.
the species affect the Darwinian fitness of each other). commitment A point in a developmental programme in which
coevolution Reciprocal evolution of interacting species that is developmental options become restricted.
driven by natural selection such that the interacting species shape community A group of organisms from different species that occur
the evolution of each other. in the same area and interact with each other.
cofactor An organic molecule or inorganic ion that is needed for comparison matrix A matrix specifying the context-independent
the catalytic activity of an enzyme. score between any two amino acids or nucleotides in two different
cognate Describes an aminoacyl-tRNA synthetase and its corre- sequences.
sponding set of tRNAs that are specific for the same amino acid. compartments Regions of a tissue in which cells do not mix across
cohesion species concept The idea that species are the most a common boundary, usually because the cells have differential
inclusive populations of individuals that have the potential for adhesive properties.
phenotypic cohesion through intrinsic cohesion mechanisms. compatible Describes fungal sexual partners that are able to
cohesion–tension theory A widely accepted mechanism for the produce a fertile cross.
ascent of water in plants. Using energy from transpiration, water is compatible solute or osmolyte Organic compound synthesized
pulled from above, creating a negative pressure which is transmitted or taken up by halophilic and halotolerant organisms to balance
all the way to the soil by the cohesion of water molecules. the osmotic strength of the medium.
cohesive ends ‘Sticky ends’, complementary single-stranded ends competence (1) The ability of cells to respond to an environmental
of a double-stranded nucleic acid that has been digested with cue in a specific manner. For example head ectoderm (but not
certain restriction enzymes. trunk ectoderm) is competent to form lens in response to a neural-
cohort study See observational study. plate-derived signal. The competence of cells is usually restricted
cold-reactive autoantibody Antibody to red blood cells that to specific time windows during development. (2) The natural
binds to the cells most efficiently at temperatures below 371C. It is development in Bacillus subtilis of the ability to take up and
principally IgM. internalize exogenous DNA from the medium.

162
Glossary

complement, complement system A group of 20 or more concealed antigens Antigens that are not normally seen by the
plasma proteins (and their associated receptors and regulatory host immune system because the host is never exposed to them.
proteins) that are involved in the elimination of extracellular For example, antigens which are part of the tick’s midgut and are
pathogens in both innate and adaptive immunity in vertebrates therefore not introduced with the salivary antigens.
(and also reported from some invertebrates). When activated by concentration gradient The difference in concentration of an ion
an infection, they can opsonize the infecting microorganism, or solute on the two sides of a membrane. If the membrane is
facilitating its phagocytosis, cause inflammation, and attack cell permeable to the solute, it tends to diffuse from the side of high
membranes, causing cell lysis. concentration to the one of lower concentration until the
complementarity-determining regions (CDRs) The hypervari- concentrations on both sides are equal.
able regions of an immunoglobulin molecule, consisting of three conceptus All products of the fertilized egg, both embryonic and
regions (loops) from the heavy chain and three from the light extraembryonic.
chain, that come together to form the antigen-binding site. conchicolous habit Of molluscs, the habit of living in shells.
complementary base pairing Specific hydrogen bonding be- concurrent asynchronous lactation Simultaneous production of
tween complementary purine and pyrimidine bases that holds the two very different types of milk from adjacent mammary glands
antiparallel strands of the DNA double-helical structure for two young of different ages.
together, and enables single-stranded RNA of complementary conditional gene knockout Inactivation of a gene in particular
sequence to be transcribed from DNA. Guanine pairs with cells and/or at a particular time through gene targeting techniques.
cytosine, and adenine pairs with thymine (in DNA) or uracil (in conditional lethal mutation A mutation in a gene essential to
RNA). Base-pairing has been exploited in the techniques of growth, in which the growth blockade can be bypassed under one
nucleic acid hybridization, PCR and sequencing methods. condition but not another. An example is a temperature-sensitive
complementation Restoration of a missing function in a mutant mutation, which restricts growth at one temperature but not at a
cell or organism, usually by introduction of a functional gene slightly lower temperature.
from another cell or organism of the same type. configuration The spatial variation that is possible in organic
complementation studies Experiments that enable the identifi- molecules, as the result of the tetravalency and the tetrahedral
cation of genes with similar or redundant function, by testing the nature of carbon atoms. Different configurations can be inter-
ability of one gene to fulfil the function of another that is absent or converted only by breaking and remaking bonds. D and L isomers
deficient. and cis and trans isomers represent different configurations.
complementation test (1) Genetic cross between a parent confinement Limitation of, for instance, motion to a specific part
heterozygous for mutation A and a parent heterozygous for of space.
mutation B in order to test whether mutations A and B are confocal microscope A microscope in which the illumination and
affecting the same gene. (2) A test in which two different nuclei or the image detection are confined to the same spot in the specimen
genomes bearing recessive mutations are associated in single cells using a spatial filter.
(heterozygotes or heterokaryons) to determine if together they conformation Different three-dimensional arrangements of a
will impart a normal phenotype. If so, they are said to molecule that can be interconverted by a simple rotation about
complement each other, and therefore the mutations lie in a single bond, without breaking any covalent bonds.
different genes. Confuciusornithids Primitive flying birds from the Late Jurassic–
earliest Cretaceous of China, including Confuciusornis sanctus
complement-like factors Circulating proteins found in some
and Changchengornis hengdaoziensis.
invertebrates that appear to be similar to the complement
congenic Describes strains of animals that are identical at all but
proteins of vertebrates.
one chromosomal region.
complete remission Total absence of symptomatic, physical and
congenic mouse strains Inbred strains of mice which are identical
laboratory evidence of a given disease.
except for a short chromosomal segment encoding a different
complex spike Action potential with a complex configuration, histocompatibility antigen.
induced in a Purkinje cell by activation via a climbing fibre. congenital heart defects (CHD) Structural heart abnormalities
complicated migraine Persisting neurological deficits that start in that are present at birth.
the aura or during the headache phase, but continue for 24 hours congestive heart failure (CHF) A cardiac disorder in which the
or more. normal function of the heart is impaired so that blood ejection is
compression A shortening of a structure that results from an decreased and blood return from the venous system is impeded.
applied force (negative linear strain). conidiogenous cell A cell that directly produces a conidium. It is
computed tomography (CT) An imaging technique in which often a specialized structure, the conidiophore.
multiple X-ray beams are used to obtain numerous projections conidiophore A specialized hypha that bears one or more
through the body. Images of slices through the body (tomograms) conidiogenous cells.
are generated by computer. Also known as CAT (computer- conidium (plural conidia) A thin-walled nonmotile deciduous
assisted tomography) and sometimes as CAT scanning. However, asexual fungal spore that is produced at the tip of a specialized
scan is a confusing word and is best avoided in the context of hypha or by fragmentation of a hypha.
imaging. conjugate base The deprotonated species remaining after an acid
computer-assisted tomography (CAT) See computed tomogra- donates a proton. The original acid and its base are known as a
phy. conjugate acid-conjugate base pair.
concatemer (1) DNA molecule that contains multiple tandem conjugate planes Those planes that are equivalent in an optical
head-to-tail repeats of a particular sequence. (2) Giant molecule system. An object placed in one plane will be imaged sharply in
consisting of multiple copies of covalently joined units. each subsequent plane of that series.

163
Glossary

conjugation (1) Close joining between two unicellular organisms convection Heat transfer between the air and an object of different
during which genetic material may be exchanged. (2) The sexual temperature.
transfer of plasmid or chromosomal DNA between bacteria that convolution A mathematical operation that extracts particular
requires direct contact between the donor and the recipient cell. features from a function or an image.
The delivery system is encoded by sex plasmids in the donor cell. cooperativity (1) In a protein with multiple binding sites, the
(3) The temporary fusion of two cells of ciliate protozoans to influence of bound ligands on the affinity of the molecule for
allow exchange of gametic nuclei. it typically results in fertiliza- binding subsequent ligands. (2) A change in enzyme activity that
tion and the formation of zygote nuclei in both participating comes from interactions between the protein subunits that help to
ciliates. stabilize a new conformation.
conjugative plasmid A self-transmissible plasmid that encodes all coordinate bond A metal–ligand bond resulting from the inter-
the functions needed for its own intercellular transmission by action of a Lewis base (the ligand) and a Lewis acid (the metal
conjugation. ion).
connective Tissue that keeps an anther together and connects it to coproantibodies Secretory antibodies that can be detected in
the filament. intestinal contents and in fresh faecal substrate.
conodonts Tiny, tooth-like fossils of calcium phosphate from the coproantigen Antigen detectable in stool.
Cambrian to the Triassic. They are thought to represent a coprolite A fossilized faecal pellet. Literally ‘dung stone’. Occa-
complex, tooth-like bony jaw apparatus that belonged to small, sionally the term is used also to describe the fossilized intestinal
eel-like animals that were probably primitive vertebrates. The contents of some creatures (notably fish).
whole conodont animal is rarely found fossilized because of its coprological examination A method of detection of infective
soft body, but conodonts themselves are common. stages of parasites in the faeces of animals or humans.
consensus sequence A theoretical sequence that is derived by coprophagy Ingestion of faeces. It is nutritionally important for
comparing a group of related sequences and designating the most some animals because it allows recovery of otherwise lost protein,
commonly occurring residue that is found at each aligned vitamins, and other fermentation products.
position.
co-protease A specific activity of the RecA nucleoprotein filament
conservation biology An academic discipline that studies biolo-
which allows it to bind to a protein and cause that protein to
gical diversity and develops methods to protect it.
cleave itself.
conserved protein sequences Amino acid-sequences that share
copulatrix See bursa.
at least 70% identity.
copy number The number of identical plasmid copies per cell or
conserved sequences Nucleic acid or amino-acid sequences are
per chromosome.
said to be conserved when they are similar in different organisms.
Conservation is considered evidence for some type of function. co-receptor In immunology, the term often refers to the CD4 and
Sequences without function usually diverge rapidly in the course CD8 molecules, which aid the binding of T-cell receptors to class
of evolution, II and class I MHC molecules, respectively.
Conservitat Lagerstätten Exceptionally well-preserved biotas coronal the plane of the body which bisects it from side to side, and
consisting of organisms that do not normally form part of the which runs at right angles to the anteroposterior, or sagittal,
rock record, principally those that have an unmineralized or plane.
poorly mineralized skeleton. corrinoid A compound resembling vitamin B12 and containing a
consortia Assemblages of different species of organisms in which porphyrin-like corrin ring displaying a central atom of cobalt.
all species derive benefit from each other (See also biofilm). cortex (1) Generally, the outer layer of an organ. (2) The outer layer
constitutive Unregulated production. of the brain, in which the neuronal cell bodies are organized in a
constitutive activity of a protein State in which a protein is in the layered manner. (3) The layer of cytoplasm immediately below
activated state in the absence of the ligand that is generally needed the plasma membrane in cells. (4) The thick peptidoglycan shell
to induce its activation. surrounding the core of a bacterial endospore.
constitutive heterochromatin Regions of highly repetitive cortical alveoli Membrane-bounded flattened vesicles or sacs
sequences of DNA that contain no genes. in the cortex or pellicle of various protozoan protists, speci-
consumer An organism that obtains nutrition by eating other fically found in dinoflagellates, ciliates and sporozoan apicom-
organisms. plexans.
contact hypersensitivity T cell-mediated hypersensitivity reac- cortical blindness Rare form of blindness arising from dysfunc-
tion produced in the skin of a sensitized individual on contact tion in the visual cortex rather than in the retina and optic
with certain chemicals. nerve.
contig A contiguous segment of DNA sequence assembled from cortical death A proposed standard for declaration of death based
partially overlapping DNA clones. on the permanent loss of brain cortical function.
continuous cell line A culture that can be passaged indefinitely cortical map The spatially orderly projection of a receptor organ
without becoming senescent. These cultures are often referred to on to the cerebral cortex.
as ‘immortalized’ cells. cortical plasticity Reversible modification of the functional
contractile vacuole A clear vacuole within a cell which swells by organization of a cortical region associated with sensory
accumulation of fluid from the cytoplasm and contracts to expel experience or sensory learning.
the fluid out of the cell. cospeciation Speciation in two or more interacting evolutionary
contralateral On the opposite side of the brain. lineages, such as host and parasite, where each speciation event in
contrast A relative difference in brightness (or in colour) between one lineage results in a matched speciation event in the other
two objects or between two areas in an object or image. lineage. The process may or may not involve coevolution.

164
Glossary

costa Slender rib-like structure subtending the undulating mem- crinoid ‘Sea lilies’, a class of echinoderms (phylum Echinodermata,
brane of typanosomatids. class Crinoidea) typically bearing an elongate stem or column
cosuppression See post-transcriptional gene silencing. that elevates a flower-like crown with feathery arms used for
Cot The initial concentration of DNA (in mole nucleotides per litre) suspension feeding. They were both abundant and diverse for
in a hybridization reaction multiplied by the duration (in seconds) several hundred million years from the Ordovician onwards.
of the incubation. They are relatively rare today and generally confined to deep
cotranslational Describes any process that acts upon a polypep- waters.
tide chain while it is being synthesized. crista The sensory organ of the semicircular canal, containing hair
cotranslational modification A modification to a polypeptide cells. It is situated in an ampullary enlargement and covered with
chain that occurs while it is being synthesized. a cupula.
cotransmitter Many neurons use more than one neurotransmitter cristae Invaginations of the inner mitochondrial membrane that
for communication with target cells, for example an amino acid form a series of projections of various shapes.
such as glutamate and a neuropeptide. In this situation each of the criterion A combination of incubation temperature and solvent
neurotransmitters is referred to as a cotransmitter. composition chosen to dissociate DNA duplexes having more
cotransport Transmembrane transport of a substrate coupled to than a specified amount of base mismatch. As the criterion
that of another substrate in the same direction. increases, the amount of base mismatch that the duplexes can
cotransporters Integral membrane proteins that mediate the have decreases.
coupled transport of ions and other substrates across a cellular critical micelle concentration (cmc) The concentration of
membrane. These proteins often use the energy gained by moving detergent at which micelles begin to form.
an ion down its electrochemical gradient to move the other ion or critical region The minimal region of a chromosome which, when
substrate up its gradient. altered, results in a specific phenotype or syndrome.
Coulomb force Force between two charges. cross transmission Transmission of an infectious agent from one
countertranscript An RNA that is encoded by the opposite strand vertebrate group to another, e.g. from reptiles to mammals.
of DNA to that encoding the mRNA for a particular gene. Not all crossing-over Exchange of chromosomal segments between
countertranscripts are antisense RNAs that regulate target homologous chromosomes during meiosis, producing new
function. combinations of alleles.
countertransport Transport of a substrate coupled to that of cross-linker Chemical that conjugates different parts of proteins
another substrate in the opposite direction. such as amino, carboxy or thiol groups, and thereby generates
coupling The process whereby osteoclast activity is linked to protein complexes.
osteoblast activity, coordinating bone resorption with bone cross-reaction Reaction of an antiserum with a molecule not
deposition. present in the immunizing preparation. It usually reflects
covalent bond The sharing of a pair of electrons between two structural similarity with the immunizing antigen.
atoms. crown gall Galls caused by the pathogenic bacterium Agrobacter-
coxa The most proximal segment in the arthropod walking leg. ium tumefaciens on dicotyledonous plants.
Coxsackie and adenovirus receptor The cell-surface receptor for crural Pertaining to the leg.
most adenovirus serotypes. cryoelectron microscopy A technique that allows macromole-
CpG island Stretches of DNA (500–1000 nucleotides) rich in cules to be visualized in their native state in a thin (~50–80 nm)
guanine plus cytosine, usually flanking housekeeping genes and layer of vitreous ice.
most tissue-specific genes. cryopreservation Preservation in a frozen state, for example in a
cranial fenestration In vertebrate evolution, upper and lower deep-freeze or a liquid-nitrogen refrigerator.
temporal windows first (fenestrae) developed just behind the cryoultramicrotome An apparatus for cutting, at low tempera-
orbit (diapsid condition), and later a window just in front of the ture, ultrathin sections of frozen material for cryoelectron
orbit also appeared (archosaur condition). These cavities then microscopy.
increased in size, making the skull much lighter. cryptic plasmid A plasmid for which there is no known phenotypic
cranial kinesis (1) The mobility of the upper jaw, or parts of it, in function but which replicates and spreads to other cells.
relation to the braincase. All birds that have been studied cryptobiosis Ability of an organism to form a resting stage in order
adequately have some degree of cranial kinesis. (2) The capacity to survive unfavourable (e.g. drought) conditions.
for movement (usually due to movable joints) between bones of cryptococcosis Invasive infection caused by the fungus Crypto-
the normally rigid skull roof. This occurs notably in bony fish, coccus neoformans.
many lizards, snakes and birds. It has also been recognized in cryptozoic Refers to the fauna that lives hidden beneath the soil/
some dinosaurs. leaf litter surface or beneath bark and other similar surfaces.
cranium The vertebrate skull minus the mandible. crystal cell Haemocyte comparable to prohaemocyte of some
CREB Cyclic adenosine monophosphate (cAMP) responsive ele- invertebrates.
ment-binding protein. A transcription factor regulated by cAMP. crystal lattice The three-dimensional distribution of points defin-
Crenarchaeota A kingdom of the Archaea that contains primarily ing the origin of each copy of the unit cell in a crystal.
hyperthermophiles. crystal unit cell The repeating unit of a crystal, which periodically
cresta Fibrillar, noncontractile structure located below the basal reproduced in three dimensions yields the crystal.
portion of the trailing flagellum of devescovinid flagellates. crystallins Major soluble proteins of the lens of the eye, which
Cretaceous period The last period of the Mesozoic era. fill the cytoplasmic space of mature lens cells. Crystallins are
cribriform plate A thin bone separating the olfactory epithelium thought to maintain the transparency and reflective index of the
from the frontal brain area. lens.

165
Glossary

crystallized song See stereotyped song. cyclin-dependent kinase inhibitor Regulatory protein of the
crystallography The study of protein structure by means of X-ray eukaryotic cell cycle that inhibits the activity of cyclin-dependent
diffraction through crystals of the protein. kinases.
ctenidium Molluscan ‘gill’, which originally only created the water cyclin-dependent kinases (CDKs) Regulatory proteins of the
current, but which is also involved in respiration in larger eukaryotic cell cycle which become activated when complexed
animals. with the appropriate cyclin.
CTL Cytotoxic T lymphocyte, the CD8 T cell type with cell-killing cyclins Regulatory proteins of the eukaryotic cell cycle whose level
capacity. Their T-cell receptors recognize peptides presented by in the cell rises and falls during a single cycle. They are involved in
major histocompatibility complex (MHC) class I molecules. the progression from one stage of the cycle to the next and there
C-type lectins Lectins that bind their carbohydrate ligands in a are several different types, acting at different stages in the cycle.
calcium-dependent fashion. Seven separate families of C-type cyclol A structure in which polypeptide chains fold into a planar
lectins have been described and all share a characteristic hexagonal array.
carbohydrate recognition domain (CRD). cyclosis Movement of subcellular components, also known as
current Flow of charged particles. Current is measured in amperes cytoplasmic streaming.
and is defined in terms of the number of elementary charges cyclostomes The most primitive class of vertebrates.
moving per second. CYP21 genes Cytochrome P450 or steroid 21-hydroxylase genes.
current-to-voltage converter An electronic device consisting of cyst A nonmotile reproductive or resting form of a cell, especially a
an operational amplifier and a resistor that measures current and protozoon, that is enclosed in a rigid secreted envelope, external
converts it to a voltage that is easily displayed and recorded. to the plasma membrane, that limits free movement of the cell. In
curve fitting An analysis method in which the correlation of data this form it is resistant to unfavourable environmental condi-
points is expressed in terms of the parameters in a mathematical tions.
function. For example, pairs of x, y points may be fit to the mathe- cystic Describes an arrested development of the ear that shows only
matical function describing a straight line, in which case the one largely undivided central lumen much like an otocyst.
parameters determined will be the slope and y intercept. In cystocyte A misnomer for coagulocyte (granulocyte).
sedimentation, the functions are derived from hydrodynamic and cytochrome bf complex A membrane-bound electron transfer
thermodynamic theory, and the parameters are those sought by the protein complex found in all oxygenic photosynthetic organisms
experimenter (e.g. s, D, Mb, etc.) that oxidizes reduced plastoquinone and reduces plastocyanin (or
cushingoid Resembling the signs and symptoms of Cushing disease: cytochrome c).
buffalo hump obesity, striations, hypertension and osteoporosis. cytochrome c A mobile haem-containing electron transport
cut and paste transposition Type of transposition in which the protein in the mitochondrial respiratory chain.
transposable element is disconnected from the flanking DNA in cytochrome oxidase The terminal protein complex of the electron
the donor site by double-strand breaks and is then joined to the transport chain, which reduces molecular oxygen to water.
target DNA. cytochrome P450 A family of haem-containing enzymes that
CVHD See graft-versus-host disease. serve as monooxygenases by activation of dioxygen. One oxygen
C-value The amount of DNA in the nonreplicated haploid genome, atom is inserted into the substrate, while the second atom is
given in picograms or base pairs. released as water. They are membrane-bound enzymes found in
C-value paradox The fact that C-values for closely related species the endoplasmic reticulum and convert non-polar chemicals to
can differ enormously and there is no close relationship between water-soluble, excretable forms.
C-value and the degree of complexity of various multicellular cytochromes Family of haem-containing electron transport
organisms. It arises from the presence of different amounts of proteins present in the respiratory and photosynthetic electron
repetitive or ‘junk’ DNA in different species. transport chains of mitochondria, chloroplasts and bacteria.
cyanobacteria (singular cyanobacterium) Ubiquitous oxygenic cytocidal Causing lysis or dissolution of a cell.
photosynthetic prokaryotes that are evolutionarily ancient and cytogenetics The study of cellular genetics at the level of the
that have maintained a significant presence on Earth in essentially chromosome.
all habitats. Previously called blue-green algae. cytokeratins Intermediate filament proteins characteristic of
cyanocyte Haemocyanin-containing haemocyte found in Porifera, epithelial cells.
Mollusca and some Arthropoda. cytokines The general term for a large group of small soluble
cycle The period of an oscillation. proteins that are secreted by cells and affect the behaviour of
cyclic AMP (cAMP) The intracellular signalling molecule cyclic other cells, for example cell proliferation, differentiation and
adenosine monophosphate, which is generated by the hydrolysis migration. They include, for example, the leukocyte interleukins
of adenosine triphosphate (ATP) by G-protein-dependent and numerous growth factors and differentiation factors.
adenylyl cyclase. Cytokines act by binding to specific receptors on target cell
cyclic AMP receptor protein (CRP) This transcription factor, surfaces. Many cytokines are secreted by immune system cells
which is triggered by cyclic AMP, activates scores of promoters during an immune response, and act locally and distantly to
and plays a major role in bacterial responses to the availability of modify the response to infection.
different sugars. At some promoters it functions to repress cytokinesis The last step of cell division, in which the cytoplasms of
transcription initiation. both daughter cells become separated. It should be distinguished
cyclic GMP (cGMP) The intracellular signalling molecule cyclic from karyokinesis, the nuclear separation during cell division.
guanosine monophosphate. It is generated by the hydrolysis of cytolytic The phase in the viral life cycle in which virions are
guanosine triphosphate (GTP) by G-protein-dependent guanylyl produced and released, destroying the host cell.
cyclase. cytomegalovirus A species-specific herpesvirus.

166
Glossary

cytomorphology The form or structure of individual cells, usually days postcoitum (dpc) The number of days elapsed since
discerned by microscopic examination. conception. It is often used as an approximate guide to stage
cytopathic effect, cytopathicity Characteristic deleterious mor- mouse conceptuses.
phological changes in host cells, including cell killing, following DDT Dichlorodiphenyltrichloroethane, a synthetic chemical pesti-
viral infection. cide.
cytophilic antibody Antibody molecules that bind to a cell, de novo biosynthesis The synthesis of molecules from the simplest
sometimes rather weakly but sometimes strongly, by regions of precursors.
the molecule other than the antigen-binding site. Thus the de novo synthetic pathway Synthesis of complex nucleotide
binding is nonspecific, i.e. not related to the presence of the molecules from simple precursors such as ammonia or amino
specific antigen that evoked production of the antibody. acids.
cytoplasm The collective name for all the components of a cell that deactivation The suppression of an activity, e.g. the return of open
are not associated with either the cell wall or the genome. In a ionic channels to the closed state.
eukaryotic cell, it comprises all organelles and material outside dead space A volume in the respiratory tract that is renewed at
the nucleus. It consists of a continuous aqueous phase (the each respiratory cycle but that does not contribute to gas
cytosol) and the organelles and inclusions suspended in it and is exchange. In mammals, the upper airways are filled with inspired
the site of most of the chemical activities of the cell. air which never reaches the alveolar space and thus constitutes a
cytoplasmic determinant An mRNA or protein which is dead space.
segregated into only one of the daughters of a mother cell and deadenylation The enzymatic removal of the poly(A) tract from
causes this cell to execute a fate different from that of its sister. the 30 region of an mRNA.
cytosine Commonly occurring pyrimidine base, one of the four death receptors A family of transmembrane receptors which
types of bases in RNA and DNA. deliver intracellular signals that culminate in proapoptotic and
antiapoptotic responses.
cytoskeleton A network of proteinaceous filaments within eukar-
yotic cells, which gives a cell its shape, is involved in motility, and death-receptor-mediated apoptosis A highly regulated form of
organizes the position of internal organelles. It is composed transmembrane signalling where the outcome may be cell death.
primarily of actin microfilaments (diameter 5 nm), intermediate decapping The enzymatic removal of the cap from the 50 end of an
filaments (diameter 10 nm), and microtubules (diameter 25 nm), mRNA.
which are associated with motor proteins such as myosin, dynein decarboxylation In C4 plants, the removal of carbon dioxide from
and kinesin. The cytoskeleton is anchored to the plasma four-carbon acids in bundle sheath cells.
membrane. decubitus ulcers Ulceration caused by prolonged pressure caused
cytosol The fluid constituents of the cytoplasm. by lying in bed for long periods.
dedifferentiation The loss of specific characteristics from a
cytostome The cell ‘mouth’ in certain protozoa, the site at
terminally differentiated cell.
which food particles are ingested and food vacuoles are
DEFC24 An ecotype of Arabidopsis thaliana.
formed.
defective interfering (DI) particles Virus particles that contain
cytotoxic lymphocyte antigen 4 (CTLA4) Cell-surface
less than a complete genome. They are dependent on the presence
molecule on activated lymphocytes, which tends to inhibit
of homologous undeleted helper genomes for their replication,
activation.
and interfere with the replication of infectious virus.
cytotoxic T cells, cytotoxic T lymphocytes (CTLs) T lympho-
defence An organism’s mechanism of reducing potential damage
cytes that can kill virus-infected cells displaying viral antigens or by another organism.
cells bearing other foreign antigens. Most cytotoxic T cells carry
defensins A family of cationic proteins that lyse pathogens by pore
the CD8 co-receptor and act against target cells that present the
formation.
specific peptide antigen in association with a major histocompat-
defensive membranes Immunologically protective membranes
ibility complex (MHC) class I molecule.
secreted by some insect parasites to avoid recognition by the
cytotoxicity The property of being able to kill cells. insect’s immune system.
deficiency a lack or defect.
definitive haematopoiesis The formation of blood cells in the
D6S45 See RD gene. fetus or adult. Erythrocytes in the adult stage express different
3D reconstruction Construction of a three-dimensional model of globin chains from those expressed during embryogenesis.
an object from several two-dimensional views of it. definitive host The host in which sexual recombination of a
D period The time between termination of DNA replication and parasite takes place.
cell division. In Escherichia coli the D period, for cells growing degree of esterification (DE) The percentage of carboxyl groups
between 20 and 60 min doubling times, is a constant of of a pectin preparation in the methyl ester form.
approximately 20 min. degree of polymerization The number of monomeric units in a
Dam methyltransferase Enzyme that transfers methyl groups to molecule of a polymer.
the adenine of GATC sites. DE See degree of esterification.
dauer larva Alternative third larval stage of Caenorhabditis dehiscence The natural release of a seed or fruit from the plant.
elegans, induced by starvation in early larval development, with dehiscent Describes a fruit that opens by splits or pores.
many morphological differences to the normal L3 larva. It is dehydroascorbic acid A reversibly oxidized form of ascorbic acid.
long-lived and resistant to stress. From the German dauer, dehydrogenase An enzyme that catalyses oxidation/reduction
enduring. reactions, usually involving NADH or NADPH.

167
Glossary

delavirdine Non-nucleoside analogue inhibitors of human dermopterans An order of placental mammals having
immunodeficiency virus 1 (HIV1). exceptional adaptations for gliding, also known as flying lemurs
delayed hypersensitivity, delayed-type hypersensitivity or colugos.
(DTH) T cell-mediated type IV hypersensitivity. An immune desaturase Enzyme that inserts a carbon–carbon double or triple
response to subcutaneous injection of the relevant antigen that bond into a molecule.
results in oedema and inflammation, which reaches a maximum desmotubule The central membranous component of plasmodes-
after 24–48 hours. It involves T cells and macrophages. mata, structures that connect the cytoplasms of plant cells. It is
deme A small group of randomly mating individuals that is more believed to be derived from endoplasmic reticulum.
or less isolated from other such groups. In some models, mating desoxyanthocyanidin, deoxyanthocyanidin Anthocyanidins
within demes occurs between genetic relatives and is not random. lacking a hydroxyl group in the 3-position.
demographic stochasticity The temporal variation in birth and desulfurylation The enzymatic removal of sulfur from sulfur-
death rates due to chance sampling of individuals, even though containing compounds.
the environment remains the same. detectors In computed tomography (CT), an array of individual
dendrite A long, branching outgrowth or extension from a neuron measuring devices that capture the photons that have passed
or nerve cell, which carries electrical signals from synapses to the through the patient. Scintillation crystals and gas-filled ioniza-
cell body (unlike an axon which carries electrical signals away tion chambers are both used.
from the cell body). A typical vertebrate neuron extends many detergents Amphiphilic molecules that lyse cell membranes
dendrites but only a single axon. and solubilize lipids and hydrophobic proteins. Examples are
dendritic Describes a cell with many long thin processes extending Nonidet P-40 and Triton X-100.
from it. determinant See antigenic determinant.
dendritic cell The professional antigen-presenting cell that takes determination In development, a stable change in cell fate.
up proteins and presents peptide antigens to T cells in conjunction detritivore An organism that feeds on dead plant and animal
with accessory molecules that stimulate T-cell activation. matter.
dendritic spine Wine glass- or mushroom-shaped protrusion from deuterostomes Animals (e.g. Echinodermata, Tunicata and
a dendrite that represents the principal site of termination of Chordata) in which the oral and anal openings are derived from
axons from excitatory neurons on interneurons, especially in the two different openings during development.
cortical regions. developmental host A host in which the parasite can complete its
dendritic transport The active transport of cytoplasmic constitu- development. Sometimes called the final host.
ents within a dendrite. diacylglycerophospholipids The most abundant class of mem-
dendrogram In phenetic (numeric) systematics, a tree-like brane lipids in all organisms (except archaea), forming the bulk of
representation of shared similarities between taxonomic the lipid bilayer. They are amphipathic phospholipids containing
units. two fatty-acyl chains (the hydrophobic portion) esterified to a
denervation Removal of nerves that innervate a muscle or other glycerol backbone, and a phosphate group linked via a
tissue. phosphodiester linkage to both the glycerol backbone and a
densitometry Technique by which the intensity of a particular polar alcohol (the hydrophilic headgroup). Examples are
chemical species is quantified using a relative measure of the light phosphatidylcholine, phosphatidylethanolamine and phospha-
absorbed. tidylinositol.
deoxyribonucleic acid (DNA) See DNA. diagenesis The physical and chemical modifications that occur in
depletion force The force that drives macromolecules together. It rocks, and fossils, after burial.
is due to entropy effects associated with the presence of other diapause (1) A regulated transient delay in embryonic develop-
macromolecules. ment, usually at the blastocyst stage. (2) A state of arrested
depolarization Cells have a transmembrane electrical potential of development in insects that is characterized by a strong
the order of –70 mV (inside negative). A change in the membrane suppression of metabolic rate.
potential towards zero is termed a depolarization. diapedesis The movement of leukocytes through the intact walls of
deposit feeding Obtaining nutrients from sediments. blood vessels.
depotentiation A decrease in synaptic strength produced by an diaspore Any part of an organism that is adapted for dispersal, e.g.
activity-dependent process, when this occurs after long-term a seed.
potentiation has previously been induced. diastole Period of relaxation of the ventricles of the heart.
depression See synaptic depression. diatoms Group of single-celled, photosynthetic organisms that
depth of field In microscopy, the axial depth within a specimen secrete outer coverings (tests) made of silica.
that appears acceptably sharp in an image. diauxy, diauxic growth The using up by bacteria of one
derivative chromosome Any structurally abnormal chromosome carbohydrate growth substrate before using another when
resulting from deletions, duplications or translocations. The growing on medium containing a mixture of substrates.
identity of the derivative is determined by the origin of the diazotrophic Capable of meeting all needs for nitrogen nutrition
centromere. by fixing elemental dinitrogen.
dermal bone Bone produced by the ossification of connective dicentric A chromosome with two distinct centromeres.
tissue of the dermis. See also endochondral bone. DIC See differential interference contrast.
dermatomyositis Progressive condition characterized by muscu- DIC See disseminated intravascular coagulation.
lar inflammation and weakness. DICOM Digital Imaging and Communications in Medicine. This is
dermatophytosis Superficial infection caused by dermatophytes the industry standard for transfer of radiological images and
(fungi parasitic upon the skin). other medical information between computers.

168
Glossary

dicotyledonous Belonging to the Dicotyledons, those angiosperm dilambdodont Describes a molar tooth cusp pattern with W-
plants whose seedlings have two cotyledons. shaped ridges.
dictyosome A name sometimes used for a Golgi stack in plant dimer A complex consisting of two similar or identical compo-
cells. See Golgi apparatus. nents.
didanosine Nucleoside analogue inhibitor of human immuno- dimerization The physical association of two (often identical)
deficiency virus (HIV) reverse transcriptase. Used as an anti-HIV proteins.
drug. dinitrogenase Bacterial enzyme that catalyses the essential reac-
diencephalon The part of the brain located immediately behind tion of nitrogen fixation, the reduction of N2. It is the product of
the cerebrum. the nifDK genes. Also known as the MoFe protein.
diether lipid A membrane lipid with two acyl (phytanyl) analogues dinitrogenase reductase Homodimeric protein that serves as the
of fatty acids attached to the polar head group by ether linkage. unique electron donor to dinitrogenase and is also involved in the
differential binding Differential binding arises from an enzymatic insertion of the iron–molybdenum cofactor into dinitrogenase. It
interaction that stabilizes one of successive stable internal states is encoded by the nifH gene. Also known as the Fe protein.
more than the other. dipicolonic acid (DPA) Pyridine-2,6-dicarboxylic acid, a major
differential centrifugation Centrifugation procedure by which component of most bacterial endospores, where it is located in the
subcellular particles are separated by virtue of their differences in core.
speed of sedimentation. diplegia See paraplegia.
differential interference contrast A special contrast method in diploid An individual or cell having two full sets (2n) of
light microscopy by which live unstained cells can be viewed. homologous chromosomes (with the exception of the sex
differentiation The structural and functional specialization of chromosomes). Somatic cells of sexually reproducing multi-
cells and tissues during development. It occurs by the gradual cellular organisms usually contain a diploid number of chromo-
maturation of cells with specialized structure and function from somes (in humans this is 46). Diploid cells thus have two copies of
unspecialized precursors as a result of changes in gene expression. each genetic locus.
Differentiation is usually accompanied by a restriction in diplospory Type of apomixis in which a diploid megaspore gives
potential for further development. rise directly to the embryo.
dipole magnet See bending magnet.
diffusion The movement of molecules from a region of high
directed transport Any type of transport that results in a linear
concentration (high chemical potential energy) to that of low
increase of the distance moved as a function of time.
concentration (low chemical potential energy). The movement
directional mutation pressure Misincorporation of a particular
itself is random and driven by thermal fluctuations.
nucleotide(s) during DNA replication. After a number of
diffusion constant Proportionality constant for the distance
generations, the genome exhibits a highly biased AT or GC bias
moved in a given time due to diffusion.
in nucleotide composition. Directional mutation pressure is one
DiGeorge syndrome (DGS) Genetic disorder with a broad
of the driving forces for genetic code change.
spectrum of symptoms such as cardiac defects, facial dysmorphia,
directional selection Selection acting against individuals at one
and absence of thymus (leading to immunodeficiency) and
extreme of a phenotypic distribution.
parathyroid glands. Also called DiGeorge anomaly.
directive interactions Morphogenetic interactions that lead to a
digestive caeca Pockets of the midgut which in some malaco- new developmental pathway of the responding cells and
stracans may fuse to form a digestive gland. restriction of their developmental options.
digital Describes processes that deal with discrete numerical disaggregatase An enzyme in the bacterium Methanosarcina
values, often as an approximation of an analogue signal. which solubilizes the cell wall so that the typical aggregated
digital mask A numerical array that is applied to an image pixel- clusters of cells are liberated as single cells.
by-pixel, resulting in a processed image. Digital masks are most disinfectant An agent, usually chemical, that is used to kill
often used for noise reduction, contrast enhancement and feature microorganisms.
extraction. disinfection The killing of harmful organisms.
digitigrade A foot posture in which the animal stands with the dismutation The conversion of two moles of aldehyde into an
digits flat on the ground and the sole or palm of the foot off the equimolar mixture of alcohol and carboxylate.
ground. disomy The presence in a cell or organism of two copies of a
digitonin-permeabilized cells Cells whose membranes have been chromosome or chromosomal segment.
made permeable to macromolecules by the detergent digitonin, disparity A measure of the morphological distance between taxa.
which tends to solubilize cholesterol in membranes and creates displacement reactions Reactions of organic compounds in
membrane pores. which a leaving group is displaced by a nucleophilic reagent,
dihedral angle Angle between the two planes containing the first with inversion of configuration.
and last three of four contiguously bonded atoms. disruptive selection Selection that acts against individuals ex-
dihydropteridine reductase (DHPR) The enzyme responsible for hibiting intermediate values of some phenotype. It results in two
the conversion of the inactive, oxidized pterin cosubstrate or more adaptive forms and eliminates intermediate forms.
quinonoid dihydrobiopterin (q-BH2) to the active, reduced form, disseminated intravascular coagulation (DIC) Abnormal over-
tetrahydrobiopterin (BH4). activity of the blood coagulation system that results in wide-
dikaryon A cell with two nuclei or a tissue composed of such cells, spread formation of microscopic clots throughout the circulatory
especially a fungal mycelium consisting of cells that each contain system, which may cause organ damage and failure. The
two nuclei, usually of unlike mating types. circulating blood is rapidly depleted of essential clotting factors
dikaryotic Describes a cell with two nuclei. so that it becomes unable to clot.

169
Glossary

dissociation constant (Kd) A measure of the rate at which two DNA helicase Protein that unwinds double-stranded DNA pro-
molecules dissociate from each other, and therefore a measure of cessively. In Escherichia coli, it is the product of the dnaB gene.
their strength of binding. For protein–DNA interactions, Kd is DNA labelling Attachment of a radioactive or other marker to a
usually expressed as the protein concentration at which there is DNA molecule so that it can be traced during subsequent
half-maximal binding to the DNA target. experiments, for example if it is being used as a hybridization
dissociation profile The fraction of duplexes remaining in a probe.
population of DNA molecules, plotted against the temperature as DNA polymerase III holoenzyme Dimeric multisubunit DNA
the temperature is progressively increased. replication enzyme consisting of two DNA polymerase III core
distance matrix A display that shows pair-wise quantitative enzymes and their b and g subunits, in which the leading and
measures of genetic distance between DNA sequences, usually lagging strand polymerases are linked together by the t dimer.
as numbers of substitutions. The holoenzyme carries out coordinated synthesis of the leading
distance methods Methods of determining the evolutionary and lagging strands.
relatedness of two homologous DNA sequences. They measure DNA polymerase DNA replication enzyme that joins deoxyribo-
the number of nucleotide substitutions per nucleotide site that nucleotides together in a template-dependent reaction to form a
have accumulated in the two sequences since they diverged. new DNA strand. It joins nucleotides together in the 50 -to-30
disulfide A chemical structure involving covalent bond formation direction only.
between two sulfur atoms. DNA primase Enzyme that synthesizes short RNA primers for
disulfide bridge The covalent linkage of the sulfur atoms in two DNA replication.
cysteine residues in a protein. This bond can tie two different DNA repair The cellular processes that result in the removal of
portions of a protein together. damaged or mismatched nucleotides from DNA and the
diterpenoid A lipid molecule with 20 carbon atoms that is formed restoration of the DNA to its previous sequence or to an intact
enzymatically by the condensation of isoprenyl units. structure.
diuretic A substance that increases the elimination of urine. DNA replication The cellular processes that result in the copying of
diurnal Having a daily rhythm with most activity during daylight both strands (template strands) of a parental, double-stranded
hours. DNA molecule into two daughter double-stranded DNA
diurnal tides A sequence of tides consisting of one high and one molecules each containing one parental strand and a comple-
low tide over approximately 24 hours and 50 minutes. mentary, newly synthesized strand.
diversity The number of taxa, such as species, that are present in DNA strand exchange A process whereby single strands of DNA
the biota under consideration. are exchanged between two homologous double-stranded DNA
divided circulation A circulation pattern in which blood is molecules.
pumped twice, once by the right side of the heart and once by
DNA strand exchange protein A protein that, upon binding to a
the left side of the heart, before it completes a full circuit of the
single-stranded DNA, catalyses the transfer of that strand to
body, i.e. through the respiratory circulation and through the
a homologous region of a double-stranded DNA molecule,
peripheral circulation.
disrupting the original duplex to produce a new heteroduplex.
dizygotic twins Twins that are derived from two different eggs and
DNA transposable element Transposable element that moves via
are thus not genetically identical.
a DNA intermediate.
DNA Deoxyribonucleic acid. The biological macromolecule that
doctrine of double effect A doctrine for those committed to the
carries the permanent store of genetic information in a cell. It is a
inviolability of specific moral laws. When the unintended effect of
polymer of the nucleotides guanine, adenine, cytosine and
an action meets the doctrine’s four conditions it is morally
thymine, and the genetic information is carried in the form of
permissible.
the sequence of nucleotides, which corresponds to a sequence of
amino acids in protein. DNA is transcribed to make messenger dolichol A polyisoprenoid lipid that acts as a carbohydrate carrier
RNA, which is used to direct the synthesis of proteins. and donor in glycosylation reactions.
DNA adduct A DNA base that has been covalently modified by dolipore A septal pore complex in certain basidiomycete yeasts in
reaction with a mutagen or carcinogen. which the pore wall is swollen into a barrel-like morphology.
DNA base composition The proportion of the total bases in a domain (1) A discrete, compactly folded, structural unit in a
DNA that are guanine (G) and cytosine (C), or thymine (T) and protein, which often also has a discrete function. A protein may
adenine (A), expressed as (G 1 C) content. Bacterial genomes consist of a single domain or of multiple similar or different
range between ~25% and 75% G 1 C. domains. (2) A region of DNA with a level of supercoiling
DNA damage A change in the structure of DNA (such as chemical different from that of the bulk DNA. (3) In taxonomy, the highest
modification of bases, or formation of a base adduct or a strand level of biological classification. The three domains of living
break) resulting from exposure to physical or chemical agents. In organisms are Bacteria, Archaea and Eukarya.
some cases, DNA damage may cause mutations or cell death. domestication The breeding of plants or animals for human use.
DNA double-strand break repair Repair of DNA double-strand dominance The situation in which an allele determines the
breaks either by homologous recombination or by nonhomolo- phenotype when it is present in only one copy.
gous end-joining. dominant allele An allele that determines the phenotype when it is
DNA–DNA hybridization In molecular systematics, a technique present with another allele (the recessive allele) in a heterozygote.
whereby single-stranded DNA from two species is associated dominant mutation A mutation that results in a phenotypic effect
(hybridized) into double-stranded hybrid duplex molecules. when in the heterozygous state. The mutant allele prevails over
Melting temperatures of DNA hybrids are used to estimate the normal allele. The phenotype of individuals heterozygous or
genealogical distances between species is being compared. homozygous for the mutation is similar.

170
Glossary

dominant-negative mutation A mutation that gives rise to an ecdysozoan An animal belonging to the clade containing arthro-
abnormal gene product that interferes with the function of the pods, onychophorans, tardigrades, nematodes, nematomorphs,
product of the normal allele when the mutant and normal allele chaetognaths and cephalorhynchs.
are present together. ecdysteroids Invertebrate steroid hormones involved in signalling
do-not-resuscitate A medical order that tells the healthcare team ecdysis or moulting.
that a particular patient should not receive certain procedures echogenicity The degree to which ultrasound is returned, by
such as cardiopulmonary resuscitation in the event that their reflection or scattering, from the region (usually of the body)
heartbeat or respiration stops. being scanned. It is indicated in the display by the level of
DOPA Dihydroxyphenylalanine, the catechol amino acid precursor brightness in the greyscale image.
of catecholamines. eclampsia Convulsions associated with the syndrome of pre-
dopamine The catecholamine precursor of noradrenaline and eclampsia.
adrenaline. It is also an important neurotransmitter in its own eclipse period The interval early after phage infection during
right. which no infectious particles can be recovered from infected
dormancy In spores and seeds, a period of minimal metabolic cells.
activity during which no further development occurs. ecological integrity The condition of an ecosystem that is largely
dorsal Towards the back of an animal. free from human interference and possesses a species composition
dorsoventral The axis between the back and underside of an and functional organization comparable to natural ecosystems in
animal, or the upper and lower surfaces of a leaf. the region.
dosage compensation The process by which the expression of ecological niche The role of an organism in a biological
genes on the X chromosome is adjusted in cells that contain more community, including where it lives, what environmental
than one X chromosome, so that levels of expression of X- conditions it requires, and how it interacts with other
chromosome genes are equalized in males and females. organisms.
double-stranded RNA-binding motif (dsRBD) An RNA-bind- ecological species concept The idea of a species as a lineage that
ing domain in proteins that consists of 65–70 amino acids and occupies an adaptive zone minimally different from that of any
interacts with double-stranded RNA only. other lineage in its range, and that evolves separately from all
Dounce homogenization Breaking open cells by shearing forces lineages outside its range.
generated in a Dounce homogenizer, a glass tube with a tight- ecomorph A morphological body design that fits a specific set of
fitting ball that acts as a teflon or glass pestle. ecological features.
downstream A way of referring to relative positions on nucleic economic development Projects that contribute financial bene-
acids. As DNA and RNA chains are synthesized in the 50 to 30 fits to people.
direction, downstream means ‘toward the 30 end’. ecosystem A community of organisms and the physical environ-
drift Differential survival and reproduction of organisms that is not ment with which they interact.
due to environmental interaction.
ecosystem goods and services The set of all economic goods (e.g.
drug resistance Ability of microorganisms to grow in the presence timber, fish) and services (e.g. pollination, flood control, climate
of a usually effective antibiotic. regulation) that are provided by functioning ecosystems and that
dry deposition The adsorption of gaseous or particular material sustain and enrich human lives and societies.
from the atmosphere on land, water or plant surfaces. economic threshold The economic limit above which it becomes
DTH See delayed-type hypersensitivity. advantageous to use a chemical such as a fertilizer, herbicide
duplex DNA Double-stranded DNA. or pesticide. For a herbicide, for example, the threshold might
duplex Double, usually refers to the double helix of DNA. be expressed as the number of weed seedlings per square
duration The actual ‘lifetime’ of a taxon, from origination to metre.
extinction. This is usually unknown. ecotype A genetically specialized population living in a particular
dwarf mutants Plants with an altered gene or genes that result in a type of habitat, for example, populations of plants living in alpine
short stature. environments. An ecotype is a subset of a more broadly
dysmetria Failure of precise reaching, as is revealed when a person distributed species.
tries to touch the tip of the nose with their fingertip with the eyes ectoderm (1) One of the three germ layers of a tribloblastic animal
closed. embryo. It consists of the outer tissue layer and gives rise to the
dyspnoea Shortness of breath. nervous system and the epidermis. (2) Epithelium covering the
outer surface of cnidarians.
ectopic Out of normal context and/or location.
EAE See experimental autoimmune encephalomyelitis. ectopic exchange Recombination between two related sequences
E box The specific DNA sequence to which basic helix-loop-helix (e.g. transposable elements) at nonhomologous positions in the
(bHLH) factors bind. genome.
E site Exit (E) site, the site on the ribosome where tRNA that has ectopic expression Misexpression of a gene with respect to space
discharged its amino acid is bound before it exits the ribosome. or time.
early genes The first viral genes transcribed, enabling nucleic acid ectotherm Animal in which body temperature principally
replication and other viral functions. reflects environmental temperature and is determined mainly
ecdysis The shedding or moulting of an old arthropod exoskeleton by the heat content of the immediate environment. See also
to reveal a new, larger one underneath. exothermy.

171
Glossary

ectothermy ‘Cold-bloodedness’; the condition in which an electrolytes Charged particles, or ions, that carry an electrical
animal does not generate the heat to warm its body by internal current when dissolved in solution.
means. electromagnetic radiation The energy emitted by an electrical or
edentulous Lacking teeth. magnetic source, and which obeys Maxwell equations.
editing In the context of translation, the ability of aminoacyl- electromagnetic waves Travelling disturbance comprising an
tRNA synthetases to prevent or correct mistakes in the electric field and a magnetic field at right angles to each other.
aminoacylation of tRNA. electron carrier A protein or other organic molecule that can
editosome A complex of proteins and/or RNAs that is assembled either accept or donate electrons in oxidation–reduction
for the purpose of editing RNA. reactions.
efavirenz Non-nucleoside analogue inhibitors of human immuno- electron transfer A set of serial redox reactions that results in
deficiency virus 1 (HIV1). transport of electrons or reducing equivalents from one molecule
effective population size The number of breeding individuals to another.
(Ne) in a randomly mating population that has the same effect on electron transport Transport of electrons via redox proteins such
genetic variability (i.e. gene diversity) as observed in the as cytochromes.
population under investigation. It reflects the strength of random electron transport chain Sequence of membrane-bound electron
genetic drift. carriers, which carry electrons from an electronegative electron
effector gene A gene that encodes a regulatory protein, either an donor to a more electropositive acceptor. In the electron
activator or a repressor, that can affect the expression of a transport chains of bacterial cell membranes, mitochondria and
reporter gene in a co-transfection assay. chloroplasts this transport is coupled to the movement of protons
effector lymphocytes Lymphocytes that can carry out their across the membrane, forming an electrochemical gradient that
functions without further differentiation and proliferation, for powers ATP synthesis.
example mature cytotoxic T cells.
electron paramagnetic resonance (EPR) A magnetic resonance
efferent Carrying away from. Efferent fibres carry signals
technique used to study the chemical nature and environment of
away from the central nervous system; efferent lymphatics
paramagnetic chemical species (those with unpaired electrons)
carry lymph away from lymph nodes and other lymphoid
such as organic radicals and certain oxidation states of transition
organs.
metals.
efferent nerve fibres Nerve fibres that conduct from the centre
electropherotype The migration pattern of RNA segments
(e.g. the spinal cord) to the periphery. The term broadly
subjected to gel electrophoresis.
corresponds to motor fibres.
electrophile An electron-deficient atom (e.g. a carbon atom with a
efferent lymphatic vessel A lymphatic vessel that carries lymph
partial positive charge, C/d1 ) or compound that tends to react
from a lymph node.
with electron-containing nucleophiles.
efferent projection The axons that project to a structure from a
source – the output. electrophilic Involving an electron-deficient centre.
egg activation A sequence of biochemical and cellular changes electrophoresis A procedure that separates a heterogeneous
that occurs after fertilization, leading to the release from meiotic population of similar molecules (e.g. DNA fragments of different
arrest, division of the fertilized egg and further embryonic sizes) on the basis of their movement in an electric field.
development. Separation is on the basis of the size and electrical charge of the
egg chamber Cluster of oocyte and associated nurse cells molecules. In gel electrophoresis, the mixture is applied to a gel
surrounded by somatically derived follicle cells, in polytrophic and an electric current passed through it.
ovaries of insects. electroporation Subjection of cells to a high-voltage electric field
eicosanoids A class of lipid signalling molecules including the to make them temporarily permeable to macromolecules such as
prostaglandins, thromboxanes and leukotrienes. DNA.
EJP See excitatory junctional potential. electroreceptors Sensory receptors for detection of weak elec-
elasmobranchs The sharks and rays. trical discharges.
elasticity The tendency of a material to return to its original shape electrostatic effects The use of complementary electrical
after an applied load is removed. charges by enzymes to stabilize the binding of substrate at
elastin Protein component of elastic connective tissue, present in an active site and to stabilize polar transition states in
artery walls, skin and intestinal walls. The elastin protein chain is reactions.
particularly flexible and will return to its original shape after electrotonic decay Passive decay of a voltage change, as a
being stretched. function of distance from the point of origin, as it travels through
electrical field The pattern of the electrical influence derived from a membrane.
the presence of an electrical potential and current. electrotonic propagation The passive transfer of electrical
electrocardiogram A recording of the waves of electrical activity potentials from one location to another in a neuron whose
associated with contractions of the heart. membrane contains no voltage-dependent conductances. The
electrochemical gradient The combination of the concen- strength of the signal gradually decays with distance from its
tration gradient and the electrical gradient (voltage source.
difference) of an ion distributed unequally across a biological elicitor Pathogen-encoded molecule that induces a plant defence
membrane. response such as phytoalexin accumulation, programmed cell
electrogenicity A property of a coupled transport process which death or synthesis of PR (pathogenesis-related) proteins. Elicitors
results in an increase in net charge on one side of the membrane. can be abiotic (e.g. ultraviolet light, low temperature) or biotic
The mammalian sodium pump is electrogenic. (e.g. microbial cell wall polymers).

172
Glossary

elicitor active site A domain of an avirulence factor of a plant embryophytes Collective term for bryophytes (hornworts, liver-
pathogenic microbe that is recognized by the protein product of worts and mosses) and vascular plants (ferns, fern allies,
the corresponding resistance gene in the plant. Mutations in gymnosperms and angiosperms). They have sexual reproduction
elicitor active sites result in a loss of recognition and induction of involving antheridia and archegonia (or their derivatives in the
resistance. case of gymnosperms and angiosperms).
ELISA Enzyme-linked immunosorbent assay. A sensitive test for emission spectrum The wavelength distribution of the light
the presence of an antigen or an antibody in a sample, which emitted by a light source.
makes use of enzyme-labelled antibodies as detectors. enantiornithomorphs Diverse group of flying Cretaceous birds
ELISPOT An adaptation of ELISA which is used, for example, to of worldwide distribution.
detect the frequency of T cells secreting a particular cytokine. encapsidation The enveloping of the viral nucleic acid in a protein
Individual cells are placed on a plastic surface coated with coat during virus replication in an infected cell.
cytokine-specific antibody and incubated. After removal of the encephalitis Inflammation of the brain.
cells, cytokines secreted by a cell and bound to the immobilized endemic (1) A species whose habitat is restricted to a particular
antibody can be detected as coloured spots by a second enzyme- region or specific locality. (2) The continuous presence of disease,
labelled anti-cytokine antibody. at a low level, in a population.
elongation The successive addition of amino acids to the growing end-joining repair Repair of double-strand breaks in DNA by
protein chain during translation of mRNA. At each round of rejoining the broken ends without the involvement of unbroken
elongation, one amino acid is added to the chain in a reaction homologous strands or extensive homologous sequences on
involving the ribosome, mRNA, the aminoacyl-tRNA, elonga- either side of the break. Addition or deletion of one or a few bases
tion factors and GTP. may occur.
elongation factors Proteins that participate in the elongation endochondral bone Bone produced by ossification of a cartilage
reaction of translation. model, also called cartilage replacement bone.
ELSI Abbreviation for ethical, legal and social issues. endochondral ossification Bone development and repair
emargination Reduction (of the skull roof in this case) by achieved through the intermediate step of formation of a cartilage
reduction of bone from the edges. scaffold.
Embden-Meyerhof-Parnas pathway A pathway that degrades endocommensal A symbiont that lives inside its host without
glucose to pyruvate through the conversion of glucose to harming it. It derives nutrition from this association while the
fructose-1,6-bisphosphate. Adenosine triphosphate (ATP) is host is neither harmed nor benefited.
generated during the process. endocrine gland A ductless gland that produces a secretion that
embedding medium A matrix for encapsulating or infiltrating a spreads or is transported in the circulating body fluids.
specimen. When solidified, this matrix acts as a support to endocytosis The uptake of extracellular molecules by animal
facilitate slicing of the specimen into thin sections. cells through the formation of membrane-bound vesicles. In
embryo (1) Generally, the stage in the development of a multi- receptor-mediated endocytosis specialized receptors bind
cellular organism from the zygote when it is not yet fully formed specific ligands and the complex is then internalized. In pinocytosis,
and capable of independent life. (2) In mammalian development, small amounts of liquid and small particles are taken up
the portion of the conceptus that will eventually give rise to the nonspecifically.
fetus, when in the gastrulation and presomite stages. endoderm (1) One of the three germ layers of a tribloblastic
embryogenesis The development of the embryo from the animal embryo. Endodermal cells move from the surface
fertilized egg. of the embryo into the interior at gastrulation, and develop
embryonic (1) Generally, pertaining to an embryo. (2) Refers into the gut and associated organs. (2) Epithelium lining
to those tissues in the mammalian conceptus that will form the the coelenteron of cnidarians. Also called the gastro-
fetus. dermis.
embryonic induction Interaction between two cell lineages endodermis Single layer of cells separating the root cortex from
required for the control of normal cytodifferentiation and the stele characterized by a Casparian band.
organogenesis. endogenous Originating within the body.
embryonic root Root structure formed during embryogenesis in endogenous retrovirus Retrovirus integrated into chromosomal
plants. DNA and transmitted in this form through the germline of a
embryo sac Female gametophyte in angiosperms. It contains an species.
egg cell, a central cell, and accessory cells (synergids and endogenous stages The stages of the life-cycle of a parasite that
antipodals). occur inside the host.
embryonic stem cell (ES cell) An undifferentiated cell type endolymph A modified extracellular fluid filling the inner ear.
isolated from the inner cell mass of blastocyst-stage mammalian Endolymph has an ionic composition similar to that of
embryos. Embryonic stem cells can be maintained indefinitely intracellular fluid (rich in potassium ions).
in culture. With the appropriate induction signals, they endonuclease Enzyme that breaks down a nucleic acid strand
can potentially differentiate into any somatic cell type of the into fragments by cleaving it at internal phosphodiester
body. bonds.
embryonic suspensor A product of the early zygotic divisions endopeptidase Enzyme that cleaves a protein chain at specific
in angiosperm plants. This structure provides positional and amino acid(s).
nutritional support for the developing embryo within the embryo endophyte A microorganism living within the tissues of a plant,
sac. usually in a symbiotic or commensal relationship.

173
Glossary

endoplasmic reticulum (ER) A specialized membrane-bounded enhancer A regulatory DNA sequence at which binding of
compartment that extends throughout much of the cytoplasm of gene regulatory proteins increases the expression of a linked
eukaryotic cells. It is involved in the maturation, modification gene.
and transport of newly synthesized secretory and plasma enhancer of PEV (E(var)) A second-site mutation that leads to
membrane proteins which are delivered into it from ribosomes enhancement of the position-effect variegation (PEV) phenotype,
on its cytosolic face. It is also a site of steroid synthesis and i.e. a mutation that results in less expression (more silencing) of a
calcium storage. euchromatic gene subject to PEV.
endoprotease Enzyme that hydrolyses internal peptide bonds in enteroblastic A type of repeated budding at the same location of
proteins. the cell surface in basidiomycete yeasts.
endoproteolytic Refers to the cleavage of internal peptide bonds enterocyte Absorptive cell of the intestinal wall.
within a protein chain. enterogenous Describes the condition in which the outer wall of a
endoreduplication Replication of DNA without nuclear division. conidium is formed by the ruptured conidiogenous cell and a new
endoribonuclease An enzyme that degrades RNA by cleaving at wall is laid down within the conidium.
internal phosphodiester bonds. enterohepatic circulation Situation in which drugs or their
endosialidase An enzyme that releases polysialic acid from metabolites are taken up by, or produced in, the liver, and
glycolipids and proteins. excreted by the liver in the bile which carries them into the gut.
endosomal latency The ratio between membrane-associated From the gut they are absorbed into the portal vein and carried
horseradish peroxidase activity and activity released due to back to the liver.
disruption of endosomal membranes. enterotoxin A toxin that affects the physiological function of the
endosomes Membrane-bounded cytoplasmic vesicles, deriving intestine.
initially from the plasma membrane, that are formed when entoparasitic Describes a parasite that lives within the body of the
extracellular material is internalized by endocytosis in animal host.
cells. They carry the endocytosed material to lysosomes for entropy A measure of the randomness in a system. The change in
degradation. entropy for a system is given by the ratio of the heat absorbed by
endosperm The storage tissue in angiosperm seeds. It derives from the system in a reversible isothermal process divided by the
a cell with two female polar nuclei, with which one male gamete absolute temperature.
nucleus has fused. envelope The envelope of lipid and protein that surrounds some
endospore (1) A spore formed within a cell. (2) A dormant virus particles. It is acquired as the virus buds from the cell
cell formed by Bacillus subtilis in response to nutrient through the plasma membrane. The envelope proteins are
limitation, and which is resistant to heat, desiccation and encoded by the virus and inserted into the cell plasma membrane
ultraviolet light. before budding.
endosternite A horizontal sheet of cartilage-like connective tissue envenoming, envenomation The effects of entry of venoms into
in the prosoma of most chelicerates that serves as an internal the living organisms.
skeleton for the attachment of muscles. environmental ethics A set of beliefs, exemplified by Aldo
endosteum A single layer of bone cells that covers the internal Leopold’s ‘and ethic’, which holds that all life has intrinsic value
surface of bone and physically separates the bone surface from and therefore the destruction of other species is immoral.
the bone marrow within. environmetrics The application of statistical theory to environ-
endosymbiont A microorganism that lives within the cells of mental data.
another organism to their mutual advantage. enzyme A protein that catalyses a specific biochemical reaction.
endosymbiosis A mutually beneficial symbiotic relationship in enzyme covalent intermediate A chemical entity that is directly
which one organism, the endosymbiont, lives inside the cells of on the pathway leading to the observed product(s) of an enzyme-
another, the host. catalysed reaction and that involves a covalent bond between a
endosymbiotic relationship See endosymbiosis. nucleophile provided by the enzyme and at least a portion of one
endothecium Cell layer comprising the anther wall. of the substrates.
endothelium The inner lining of blood and lymph vessels, enzyme crystal structure The atomic structure of an enzyme
formed of a single epithelial layer of flattened cells. It separates molecule deduced from the pattern of diffracted X-rays when an
the blood from the outer layers of the vessel wall, or X-ray beam is passed through a crystal of the enzyme protein.
subendothelium. enzyme inhibitor Any substance that decreases the catalytic
endotherm Animal whose body temperature is independent of activity of an enzyme, usually by binding to it.
environmental temperature and remains fairly constant, and is enzyme-linked immunosorbent assay See ELISA.
determined mainly by the balance of heat generated by enzyme repression Inhibition of the synthesis of an enzyme by a
metabolism and heat lost through thermal conductance. specific repressor substance, often the end product of the
endothermy Regulation of body temperature by the generation of pathway.
heat inside the body as the result of metabolic processes. enzyme substrate A reactant that binds to an enzyme and, while
endplate A specialized synapse connecting motor nerve endings bound, participates in the reaction catalysed by the enzyme.
and striated muscle cells. Also called a neuromuscular junction or enzyme–substrate complex See Michaelis complex.
motor endplate. epidermis The outermost layer of cells on the body of an animal or
energized membrane A membrane across which an electrical plant. In vertebrate skin, it is the outer layer of keratinocytes,
potential exists. separated by a basal layer from the dermis.

174
Glossary

Eph receptors Receptor tyrosine kinases that are activated by epithelial–mesenchymal interactions Morphogenetic or induc-
ephrins. They fall into two classes: EphA receptors bind ephrin-A tive interactions between an epithelium and its mesenchymal
ligands; EphB receptors bind ephrin-B ligands. stroma, e.g. in parenchymal organs.
ephrins Cell-surface adhesion molecules involved in axon gui- epithelium Sheet of cells that forms the linings of body cavities and
dance and other aspects of neural development. They fall into two the glandular structures of organs. Epithelia are formed of cells
classes: the ephrin-A family are tethered to the plasma membrane tightly bound together and polarized, with distinct apical and
by glycosylphosphatidylinositol anchors; the ephrin-B family are basal faces. The small portion of an antigen that provides the
transmembrane proteins. actual binding site for an antibody or (after processing of the
epibiont Organism using another organism as a substrate to live on. antigen) a T-cell receptor. A single antigen may have more than
epicuticle (1) The outermost waxy covering of the arthropod one epitope. Also known as an antigenic determinant.
cuticle. (2) The outermost body layer of horse-hair worms epitope The small portion of an antigen that provides the actual
(nematomorphs), often sculptured and containing bristles and binding site for an antibody or (after processing of the antigen) a
warts. T-cell receptor. A single antigen may have more than one epitope.
epidemic Large-scale outbreak of an infectious disease that Also known as an antigenic determinant.
spreads rapidly through the population. epitope tagging The addition of a well-characterized epitope
epidermal basal cells The proliferating undifferentiated stem cells from a known antigen (such as influenza virus haemagglutinin) to
in the basal layer of the epidermis, from which the keratinocytes another protein to make it readily detectable by antibodies
of the skin are derived. specific for the epitope.
epifluorescence Microscopical technique that uses an ultraviolet epizootic A large-scale outbreak of an infectious disease in
filter to detect autofluorescence. animals, originating from a common source.
epigenetic development Those aspects of development that are EPR See electron paramagnetic resonance.
not directly programmed by gene action. EPS (1) Extracellular polymeric substances. (2) See exopolysac-
charide.
epigenetic inheritance The transmission of a stable pattern of
EPSP See excitatory postsynaptic potential.
gene expression that is due to reversible modifications of DNA or
Epstein–Barr virus A virus of the herpesvirus family that infects B
chromatin, such as DNA methylation or histone acetylation.
lymphocytes and causes infectious mononucleosis and is also
epigenetic regulation Process by which stable modifications of
implicated in the development of certain cancers.
gene function occur (heritable through mitosis) that are not due
equatorial and axial bonds A bond to a tetrahedral atom in a six-
to changes in the base sequence of the DNA, but reflect other
membered ring system, as in glucopyranose, is said to be
changes in the chromatin. X-chromosome inactivation in female
equatorial if it makes a small angle to the plane of the ring and
mammals is an example.
axial if it makes a large angle to the plane of the ring.
epimastigote A developmental stage of trypanosomatids in which
equilibrium (1) The condition that prevails when a forward and a
the kinetoplast is located in the anterior region of the cell, in front
reverse reaction proceed with equal rates. (2) A state of a
of the nucleus. In this form the flagellum emerges laterally,
macromolecule–ligand system in which concentrations of all
forming a short flagellar pocket.
reactants remain constant (time change of any concentration
epimerase An enzyme that will racemize a single chiral centre in a C is dC/d, t 5 0). Generally, such a state is achieved in an infinite
molecule that contains more than one such centre.
time.
epimorphic regeneration Regrowth of amputated structures equilibrium centrifugation Technique by which particles are
from an anatomically complex stump. This may involve either separated by centrifugation by virtue of their different densities in
replacement of parts of appendages or regeneration of fractions a density gradient.
of organisms into new complete individuals after their bisection. equivalent Cot (Ecot) The Cot value with a correction applied for
epiphyte An organism growing on the surface of a plant. salt concentration if the solvent is other than 0.12 mole per litre
epipod A lateral process from the base (protopod) of the sodium phosphate buffer.
arthropod limb. ER See endoplasmic reticulum.
episome A plasmid that can exist either as a circular extra- erectile tissue Tissue that becomes firm or rigid due to increased
chromosomal DNA or integrated into the cell chromosome. blood flow, usually during sexual arousal.
The terms ‘plasmid’ and ‘episome’ are sometimes used inter- ergosterol A fungal-specific membrane lipid targeted by all three
changeably. classes of current antifungal therapies.
epistasis Alteration of the expression of one gene by effects at erratics Glacially transported boulders, deposited far from their
another gene locus. This is how different genes interact in the geologic source, often with a mineral composition different from
development of behaviour, physiology, and morphology. that of their resting place.
epistatic fitness interactions The fitness of a genotype is erythroblastosis The flooding of the circulation with erythro-
determined by allele combinations at multiple loci in a blasts (nucleated red cells), which occurs when haemolysis is so
nonadditive manner. severe that blood production is stimulated in sites other than the
epistome Lip along the inner tentacle row in bryozoans, covering bone marrow.
the mouth from the dorsal side. erythrocyte Red blood cell.
epithelial cell (1) Cell that forms part of an epithelium. (2) Large erythropoiesis The generation of red blood cells from undiffer-
myoepithelial cell of either ectodermal or endodermal origin in entiated precursors in bone marrow.
hydrozoans. Epithelial cells differentiate progressively as they EST See expressed sequence tag.
migrate towards the extremities, thus becoming head- or foot- electrospray ionization mass spectrometry See ESI-MS.
specific. espundia Mucocutaneous leishmaniasis.

175
Glossary

essentialism Belief in essences, or timeless and unchanging excision repair A type of DNA-repair process in which a single-
patterns, often associated with thinking in stereotypes and de- stranded tract of DNA containing damaged or mismatched bases
emphasis of diversity. Also known as typology. is excised from the DNA helix and replaced with new bases.
estuarine Pertaining to an estuary where tidal water meets a river excitable media Materials that actively respond to their environ-
current. ments by virtue of intrinsic chemical or electrical properties. All
E-tests A commercial system for estimating minimum inhibitory living tissue and cell aggregates fall into this category, and some
concentrations of antimicrobial agents. nonliving excitable media have also been described.
ethylene Gaseous plant hormone involved in regulating plant excitation (1) Of a neuron or other excitable cell such as muscle,
responses to stress, including mechanical damage. the induction of an action potential by stimuli that produce a
euchromatin Whole chromosomes or chromosomal segments that sufficiently large depolarization of the cell membrane. (2) The
show normal staining properties, the normal cycle of chromo- mechanism by which the initial signal from the chemotactic
some condensation and decondensation and DNA replication, receptor is transduced to the bacterial flagellar motor and affects
and may be genetically active. the direction of flagellar rotation.
eugenics Practices that attempt to control human biological excitatory signal See excitation.
heredity. excitation–contraction coupling The process by which the
eukaryotes Organisms of the Domain Eukarya: protists, animals electrical events associated with cell membrane depolarization
and plants. They may be multicellular or unicellular and have in muscle cells are coupled to the activation of contractile proteins
cells with a complex organization, distinguished from the cells of to initiate muscle contraction.
prokaryotes by a distinct nucleus containing the DNA in the excitatory Describes an influence tending to depolarize the
form of chromosomes, and a cytoplasm containing membrane- membrane potential of a neuron towards its action potential
bounded organelles of different functions such as mitochondria, threshold.
endoplasmic reticulum, Golgi bodies and (in plants) chloroplasts. excitatory junctional potential (EJP) The postsynaptic response
eukaryotic Pertaining to eukaryotes or their cells. of a muscle cell to activity in an excitatory motor neuron.
eukaryotic cell A cell of a eukaryote, distinguished from prokary- excitatory neurotransmitter A chemical released from nerve
otic cells by their generally larger size and complex organization, terminals that produces excitation of the target neuron.
with the genetic material enclosed in a membrane-bounded excitatory postsynaptic potential (EPSP) The depolarization of
nucleus, and a cytoplasm containing many different types of the membrane of a postsynaptic cell which brings membrane
membrane-bounded organelles. potential nearer the threshold at which an action potential will
euploid Having the normal number of each chromosome. be generated. It occurs in response to an excitatory stimulus
(neurotransmitter) from the presynaptic cell.
Europa A moon of Jupiter which may harbour a water ocean
excitotoxicity Neuronal toxicity induced by overstimulation by
beneath its icy carapace.
glutamate or related amino acids. It is due to overstimulation of
Euryarchaeota A kingdom of Archaea consisting of methanogens,
the ionotropic receptors for these molecules and the resultant ion
extreme halophiles, hyperthermophilic sulfate reducers, Thermo-
movements across the neuronal membrane.
plasma and the Thermococcus–Pyrococcus group.
excluded volume The volume around a three-dimensional object
eurytopic Adapted to a broad range of environments and so
that is inaccessible to other three-dimensional objects.
having a broad ecological range.
excrescences DNA extensions or protuberances of the bacterial
eustatic sea level An estimate of average global sea level after
nucleoid that reach far into the cytoplasm towards the ribosomes.
correcting for local variations in land surface subsidence and/or
excretory antigen Antigens excreted by an organism.
emergence.
exfoliation The constant shedding of cells from the uppermost
eutely A constancy of cell number in particular tissues or organ layers of squamous epithelium.
systems. It is common when structures are secondarily simplified exobiology Study of the origins, early evolution and distribution
(e.g. through miniaturization). of life in the Universe.
eutherian mammals, Eutheria Subclass of mammals in which the exocrine gland A gland that has a duct and produces a secretion
embryo is nourished through the placental circulation (cf. that passes out of the gland onto a body surface (external or
marsupial mammals). internal), e.g. salivary gland.
euthermy Regulation of body temperature in the range 30–451C, exocytosis The fusion of an intracellular vesicle with the cell
generally by endothermic processes. membrane and discharge of its contents to the exterior of the cell.
eutrophic Describes a nutrient-rich aqueous environment. exodermis Hypodermal cells of plant roots having a Casparian
E(var) See enhancer of PEV. band.
evaporite mineral A mineral deposited from a saline solution as a exogenous stage The stage in the life-cycle of a parasite that is
result of concentration through evaporative water loss. released outside the host to the environment.
e-vector orientation Sunlight is linearly polarized, i.e. it oscillates exon A sequence within a gene or primary RNA transcript that is
in a preferred orientation perpendicular to the direction of retained following the splicing reaction and thus represented in
propagation. Many animals can sense this orientation. the final mature RNA product. Exons usually, but not always,
evolutionary species concept A lineage evolving separately and represent protein-coding sequences.
temporally from others and with its own unitary evolutionary exon splicing Processing of a primary RNA transcript that
role and tendencies. removes intervening (intron) sequences and joins the remaining
EXAFS See extended X-ray absorption fine structure spectroscopy. exon sequences together to form the functional RNA.
excess mortality An increase in the death rate that occurs during exonuclease Enzyme that degrades a nucleic acid from the ends,
influenza pandemics and many epidemics. cleaving off one nucleotide at a time.

176
Glossary

exopeptidase Enzyme that hydrolyses one or two amino acids extracellular signals Molecules, including metabolic intermedi-
from either end of a peptide. ates, proteins and peptides, that can be exchanged between cells as
exoplanet A planet beyond the Solar System. a means of communication.
exopolysaccharide (EPS) Polysaccharide found outside the extracytoplasmic location Location of a parasite inside the cell
microbial cell wall. but without contact with the cell cytoplasm.
exoribonuclease An enzyme that degrades an RNA molecule by extraembryonic Refers to those tissues in the conceptus required
removal of successive nucleotides from either the 50 or 30 end. for implantation and maintenance of the conceptus during
exoskeleton The hard external supporting skeleton of insects and gestation, but which will be shed at birth (e.g. placenta, yolk sac,
other arthropods. It is secreted by epidermal cells. amnion).
exosporium The loosely fitting envelope of complex chemical extrarenal transplants Transplants other than kidney trans-
composition surrounding the outer coat of a bacterial plants, including heart, lungs, liver, pancreas and intestinal
endospore. transplants.
exothermy Regulation of body temperature by variation in the extrasporogonic forms Vegetative stages of a parasite that
flow of heat into the body from outside, generally by behavioural multiply in tissue locations different from the site of sporulation.
adjustments. extravasation Migration of white blood cells from the blood-
expansin An enzyme whose activity is to break hydrogen bonds. It stream into a tissue.
is thought to act primarily to break hydrogen bonds between extreme halophiles Archaea that can only live in the presence of
cellulose and molecules bound to the surface of cellulose in plant high concentrations of sodium chloride (usually more than 10%).
cell walls. extremely thermophilic sulfur metabolizers Archaea that live
experimental autoimmune encephalomyelitis (EAE) An auto- between 601C and 1131C and gain energy by the oxidation or
immune disorder of the central nervous system experimentally reduction of sulfur compounds or by anaerobic sulfate respira-
induced in mice by injection of myelin components. It serves as a tion.
model for human multiple sclerosis. extremophile An organism that exists under extreme conditions
explant Tissue removed from an organism and transferred to an of temperature, salt and/or pressure, e.g. in deep oceanic
artificial nutrient medium for growth. vents.
expressed sequence tag (EST) A clone from a cDNA library for extrinsic allergic alveolitis A type III immune complex disease
which a partial sequence has been generated. produced by the inhalation of soluble antigens.
expression system A cell culture system into which a foreign DNA exuviae The cast-off fragments of a moulted arthropod exo-
can be introduced as part of a specialized vector that enables it to skeleton.
be expressed as protein.
ex vivo Describes a type of therapy in which cells extracted from a
expression vector In recombinant DNA work, a type of vector patient are subjected to treatment and then reintroduced into the
that contains promoters and other regulatory sequences that patient.
enable a protein-coding DNA inserted into the vector to be
ex vivo gene transfer Gene transfer into cells preceding the cells’
transcribed and translated if introduced into the appropriate
implantation into the body.
cells.
extant Living, not extinct.
extender unit See starter and extender units.
extended X-ray absorption fine structure spectroscopy (EX-
AFS) An X-ray spectroscopic technique that can reveal short- F chlorosomes Chlorosomes found in the nonsulfur filamentous
range distances (less than approximately 0.4 nm) between a metal bacterial family Chloroflexiaceae.
atom and neighbouring metal and ligand atoms. Unlike X-ray F factor See F plasmid.
diffraction methods, EXAFS can be performed on non-crystal- F pilus Protein filament on the bacterial surface through which
line preparations. DNA is passed from one bacterium (the ‘male’) to another (the
extensibility A quantitative strain of a cell wall as a result of ‘female’).
the stress induced by the osmotic pressure above the yield F plasmid A sex plasmid of the bacterium Escherichia coli, which
threshold. enables conjugation and its own transfer by encoding formation
external sense organ Sensory structures that originate in the of the F pili.
periphery, with axons that project into the central nervous F0 F-prime, an F plasmid that carries a stretch of bacterial
system. They serve mechano- and chemoreceptor functions. chromosomal DNA, and can thus transfer chromosomal genes
external state Free enzyme plus free ligand. from one cell to another.
extinction The disappearance of a species, or other larger group of F1F0 ATPase The protein complex located in the inner mitochon-
plants or animals. drial membrane that synthesizes ATP from ADP and PI. The
extracellular matrix A material composed mainly of protein and energy for the reaction comes from the flow of protons through
proteoglycan that is secreted by cells and fills the extracellular the protein complex.
spaces in tissues. Fab fragment A fragment of an antibody molecule that contains
extracellular pathogens Microorganisms that replicate and/or one of the antigen-binding sites. It consists of one entire light
persist on surfaces or in the extracellular spaces of tissues. chain and the V and CH1 domains of the paired heavy chain and is
extracellular polymeric substances (EPS) Polysaccharide and obtained by papain digestion.
proteinaceous material secreted outside of the diatom frustule facies A particular kind of sedimentary environment, e.g. black
that takes the shape of stalks, tubes, films and pads. shale, shallow marine carbonate.

177
Glossary

facilitation The fastest phases of enhanced transmission in short- febrile neutropenia A clinical condition with severe neutro-
term synaptic plasticity, consisting of two components with time phil deficiency (neutropenia) (usually less than 0.5  109
constants of tens and hundreds of milliseconds. neutrophils per litre) and fever (usually temperature greater than
FACS See fluorescence-activated cell sorter. 38.51C).
factor A general term for a molecule (almost always a protein) or feral Describes an animal that is wild and untamed, not domes-
complex of molecules, that carries out a particular function or is ticated.
involved in a particular reaction. It is used especially when a fermentation A type of metabolism in which glucose (or other
molecule is identified only by its function and has not yet substrate) is partially oxidized using internal electron acceptors,
been chemically characterized. The names of many proteins, such e.g. pyruvate.
as epidermal growth factor, factor VIII, elongation factor, ferredoxin A soluble FeS protein located in the stroma of
initiation factor, nerve growth factor, arise historically from this chloroplasts that accepts electrons from photosystem I. Ferre-
usage. doxin serves as a branch point in photosynthetic electron
factor VIII Blood coagulation factor whose lack of activity in transport.
plasma causes haemophilia A. fertility An individual’s ability to produce offspring, measured
factor IX Blood coagulation factor whose lack of activity in plasma in population terms as the average number of offspring per
causes haemophilia B. parent.
facultative heterotrophs Organisms that are able to grow as fertilization The fusion of the cells and nuclei of two gametes of
chemolithoautotrophs as well as heterotrophs. opposite sex to form a zygote.
falsifiability An indicator of the scientific status of a theory, in that fetoscopy Direct visualization of the fetus through an endoscope
it is in principle open to disproof. A good scientific hypothesis will during pregnancy.
make predictions about the world that expose it to refutation by fetus In mammalian development, the embryonic portion of the
empirical evidence. conceptus once the somites have formed and beyond.
FAOS Fluorescence-assisted organelle sorting. A method of FGF, Fgf, fgf Gene symbols for fibroblast growth factor genes.
purification by which fluorescently-labelled organelles, such as FGF See fibroblast growth factors.
phagosomes containing microorganisms that express the green FH Bacteriophage of the archaeon Halobacterium salinarium.
fluorescent protein (GFP), can be isolated from other organelles fibrinogen Plasma protein that is converted to insoluble fibrin
using a fluorescence-activated cell sorter. through the proteolytic action of thrombin during the blood
clotting reaction.
Farenholz’s rule ‘Parasite phylogeny mirrors host phylogeny’.
fibroblast Cell type present in the dermis and in the connective
fasciculation The growth of nerve cell axons side-by-side to form a
tissues of other organs. It produces most of the extracellular
bundle.
matrix proteins.
FASTA format A standard format for nucleic acid or protein
fibroblast growth factors A family of signalling proteins, many of
sequences. The first line begins with a ‘greater than’ sign (4)
which are secreted and bind to extracellular matrix. They have a
followed by a short description of the sequence. The subsequent
variety of roles in embryonic development and in the adult.
lines consist of strings of one-letter amino acid or nucleic acid
fibrolamellar bone Bone tissue initially formed as a finely
codes.
cancellous framework of woven bone, which is later compacted
fate map A definition of regions in the early embryo that will
by lamellar bone that forms around blood vessels and results in
normally give rise to a particular tissue in the adult.
primary osteons.
fat-soluble vitamins Vitamins A, D, E and K, a group of fictive feeding A rhythmic neuronal firing pattern underlying the
terpenoids composed of an activated five-carbon isopentenoid generation of rhythmic feeding movements.
unit. filopodia (singular filopodium) Fine actin-based finger-like
fatty acid The carboxylic acid of a (sometimes modified) hydro- protrusions that extend and retract at the leading edge of
carbon chain. In cells, they are used as energy sources and as migrating cells. Also known as microspikes.
building blocks for more complex molecules such as membrane fimbriae (singular fimbria) Nonflagellar filamentous structures
phospholipids, fats and oils. on the surface of bacteria. Also called pili.
fauces The passage from the mouth to the pharynx. final host The host in which the sexual stages of a parasite occur.
Fc receptor Cell-surface receptor on phagocytes and other Also called the definitive host.
cells that binds the constant part (Fc region) of an fine needle aspiration The sampling of tissue or cells from the
antibody when it is complexed with its antigen. There are body by insertion of a small hollow needle, such as that used to
different Fc receptors specific for the different classes of take a blood sample, and application of negative pressure via an
antibody. attached syringe.
Fc That part of an antibody molecule comprising most of the finite cell line A cell culture that can be maintained by serial
constant region, including the binding sites for complement and passage but has a characteristic finite limit on the number of
for receptors (Fc receptors) on various types of white blood cell. It population doublings it can achieve before entering a state of
is responsible for antibody effector functions and has no antigen- senescence.
recognition capability. FIS A bacterial histone-like protein.
Fc/epsilonRI High-affinity receptor for the Fc portion of IgE, Fisher runaway process The rapid coevolution of mating
present on mast cells and basophils. preferences and secondary sexual traits.
Fc/epsilonRII A C-type lectin which is a low-affinity receptor for FISH See fluorescence in situ hybridization.
IgE, present on B cells. fistula An unnatural connection between an organ and the body
febrile With an abnormally high body temperature. surface or between organs.

178
Glossary

fitness In the Darwinian sense, the probability that an individual fluorescence-activated cell sorter (FACS) An opto-electronic
organism will survive and produce viable offspring, relative to the device used to detect and sort different types of cells from a
survival and reproductive success of other individuals in the mixture by detecting fluorescently labelled antibody bound to
population. It is thus a measure of the relative adaptive value of a their surface antigens. Antibodies specific for different antigens
given genotype in comparison to other genotypes in the are labelled with different fluorescent tags, enabling the detection
population. It can give a quantitative assessment of an organism’s and quantitation of particular cell types.
ability to pass on its own genes, in itself and its relatives fluorescence-assisted organelle sorting See FAOS.
(offspring, siblings, etc.), to future generations of the population fluorescence in situ hybridization (FISH) A technique for
of which it is a member. It is measured as the relative contribution identifying chromosomes or genes on chromosomes by the use
of an individual’s genotype to the next generation relative to the of fluorescently labelled DNA probes. The probes are hybridized
contributions of other genotypes. to chromosomes or nuclei immobilized on glass slides to
fixation (1) In genetics, the case when one allele comes to determine visually the presence or absence of a particular DNA
predominate in a population such that all individuals possess sequence.
that allele. (2) Stabilization of biological specimens for micro- fluorescence resonance energy transfer (FRET) Technique for
scopic examination in a form that retains as much as possible of investigating molecular structure and function which depends on
their natural structure. Chemical treatment converts macro- the fact that some molecular groups begin to fluoresce when they
molecules to insoluble derivatives. come sufficiently close together.
fixation index An index of genetic variation between populations, fluorescent dyes Dyes that emit fluorescent light of various
expressed in terms of gene frequency variability between colours when viewed under specially equipped light microscopes
populations, standardized in relation to the maximum possible with near-blue ultraviolet light and appropriate filters. They are
variability. used to stain cells and to label antibodies and DNA probes.
fixed lineage A stereotyped sequence of cell divisions that usually fluorophore A chemical group that fluoresces strongly when
generates a defined array of cell types. suitably illuminated, and which can be observed using a
flagellum (plural flagella) (1) A motile structure on the surface of fluorescence microscope.
some prokaryotes, whose rotation propels the microorganism flush The appearance of mushrooms at intervals (rhythmic cycle)
through an aqueous milieu. It consists of a whip-like filament for harvesting.
attached to a basal body embedded in the bacterial membrane. flux (1) The number of objects passing through a plane of known
(2) A superficially similar motile structure on eukaryotic cells, area in a unit of time. The usual units of flux are moles (or grams)
which has a different microtubule-based, molecular structure and per cm2 per second. (2) Reaction rate.
mechanism. fMet-tRNAfMet Initiator tRNA charged with formyl-methionine.
flip-flop The movement of a phospholipid from one leaflet of a FMO protein Protein responsible for the attachment of S chloro-
bilayer membrane to the other. somes to the plasma membrane.
flocculate A complex of organic and inorganic materials which FNR An enzyme located in the chloroplast stroma that oxidizes
settles out of the water column. ferredoxin and reduces NADP1 .
flood basalt province Vast province of flat-lying lava flows FNR protein A homologue of the cyclic AMP receptor protein that
that often contains up to several million cubic kilometres of is activated in the absence of oxygen. It is a bacterial
lava. More than 10 such provinces are known and they constitute transcriptional activator with an essential role in the induction
by far the most spectacular manifestation of volcanic activity on of gene expression during adaptation to anaerobic growth
earth. conditions.
floor plate The ventral midline region of the epithelium of the early focal adhesion sites Sites at which cultured eukaryotic cells make
vertebrate neural tube. contact with their substratum. Focal adhesion sites contain
floral meristem identity genes A group of genes that specify the clusters of integrins, with their extracellular parts bound to
identity of the floral meristem in Arabidopsis. extracellular matrix proteins and their intracellular parts
floral organ identity genes Genes that determine the identity of attached to actin filaments.
individual floral organs such as sepals or petals in Arabidopsis. folding of proteins The process by which a polypeptide chain
flow cytometry Photoelectronic technique for the counting and adopts the precise three-dimensional structural conformation
sorting of individual cells based on detection of granularity or that is of lowest free energy.
fluorescent labelling. folding pattern The unique three-dimensional spatial arrange-
flow cytometry Photoelectronic technique for the counting and ment of atoms in a protein molecule that has adopted its final
sorting of individual cells based on light scattering or the conformation.
detection of fluorescence. The cells pass in front of a laser beam follicle In botany, a single-cavity, multi-seeded fruit that opens by
and both scattered light (a function of size) and fluorescence can splitting.
be measured. footprint Of a transposable element, the characteristic donor site
FLT-3 ligand A growth factor for stem cells in vivo. It is a ligand for sequence that is left after transposition.
an FMS-like receptor tyrosine kinase. foramen magnum The opening in the base of the cranium of the
fluctuating asymmetry Morphological asymmetry in bilaterally skull that connects the posterior fossa and the cervical spinal canal.
symmetric organs which does not show a bias toward either foraminifera Small benthic or planktonic marine protozoans
side. (phylum Sarcodina, class Rhizopoda, order Foraminiferida) that
fluidity The degree of motion within the lipid components of the secrete tubular, commonly chambered, tests of agglutinated
bilayer membrane. It is enhanced by the presence of double bonds sediment grains or, more commonly of calcite. They are common
in the fatty acyl chains and by shortening of the acyl chain length. and important microfossils.

179
Glossary

force An influence on a body that causes it to accelerate (product of fructan A polysaccharide composed primarily of D-fructosyl units
mass and acceleration). in the furanose ring form.
foregut The oesophagus and stomach. fruiting bodies (1) The sexual or asexual spore-bearing structures
foreign antigen Any nonself tissue or substance whose presence of fungi and similar organisms. They include the ‘mushrooms’
triggers an immune reaction in the host. and ‘toadstools’ (basidiocarps) of basidiomycetes and the
forespore Cell formed after asymmetric division early in sporula- ascocarps of ascomycetes. (2) The multicellular spore-bearing
tion that is destined to become the mature spore. structures of slime moulds and myxobacteria.
fork arrest In DNA replication, the arrest (or pausing) of a fruiting body development The aggregation and coordination of
replication fork at a defined location on the chromosome. hundreds of thousands of myxobacteria to form a compact
formation In geology, a mappable rock unit with a common multicellular structure that contains myxospores.
character. frustule The silica-containing coat of diatoms which consists of
fossorial Adapted for burrowing or digging or for a subterranean two overlapping halves, the epivalve and hypovalve.
life.
FST The measure of the fraction of the total genetic variation at a
founder cell (1) Cell from which a structure or tissue originates.
locus that is among populations. It measures how genetically
(2) Early embryonic cell in Caenorhabditis elegans whose fate has
different one deme in a metapopulation is from another, when
been specified as to the division patterns it follows and the type of
averaged across loci.
cells it produces.
fucoxanthin Accessory pigment (carotenoid) in many golden and
founder event The establishment of a new population by a
brownish algae.
relatively small number of individuals.
fourfold degenerate site Any nucleotide can occupy this site in a functional food A food that provides health benefits beyond those
codon and the codon will encode the same amino acid. of basic nutrition.
four-helix bundles A protein folding pattern in which four functional genomics Determination of the function of genes that
segments of helical structure are packed together so that have been identified via genome sequencing programmes.
hydrophobic sides of the helices are inside. functional imaging The formation of an image containing
Fourier analysis A technique by which a complex mathematical information relating to physiological function.
function (e.g. a spectrum) is approximated as the sum of a series of functional mRNA half-life The length of time an mRNA
sine waves. remains sufficiently intact to allow the synthesis of a full-length
fovea A depressed region of the retina that, because of its high protein.
density of cone receptors, allows maximum visual acuity. fusion partner A protein molecule that, when combined with
foveate retina A retina with a circumscribed region of highest another, will introduce a new property to that molecule such as
acuity, the fovea. affinity for a matrix or greater stability.
frameshift A mutation that results in the loss of the correct reading fusion protein (1) A protein on the surface of a virus particle
frame. responsible for fusion of the virus envelope with cell membranes.
free energy The combination of energy, entropy, and work (2) Novel protein that is encoded by two coding sequences from
terms, the sign of which determines in which direction a given different proteins that have been joined together.
reaction will proceed. It is the energy available too do useful futile cycle A set of enzyme-catalysed cyclic reactions that
work. consumes energy (mostly adenosine triphosphate) in an appar-
free radical An atom or group of atoms possessing an unpaired ently useless process. When its role in regulation is established, it
electron, which usually makes it highly reactive. is then known as a ‘substrate cycle’.
free-flow electrophoresis Injection of a sample into a continuous
flow of buffer between two electrodes. Deflection of material is
dependent on charge and flow time (distance).
freezing point Temperature of a solution at which an ice crystal
placed in the solution begins to grow. G 1 C content The mol% of the bases guanine and cytosine in the
frequency facilitation See facilitation. chromosomal DNA. The range in known genomes is between
frequency-dependent selection Selection that increases or ~22–74 mol%.
decreases depending on the relative frequency of an allele in a G proteins Guanine-nucleotide-binding proteins. A large family of
population. proteins that act as molecular switches in intracellular signalling
Fresnel reflection coefficient A number that expresses the pathways. They exist in an active state when GTP is bound and in
fraction of the intensity of unpolarized light partially reflected an inactive, conformationally altered, state when the bound GTP
at a surface boundary between two transparent substances with is hydrolysed to GDP by the G protein’s intrinsic GTPase
different refractive indices. activity. The term G protein itself is usually reserved for the
FRET See fluorescence resonance energy transfer. heterotrimeric G proteins that transmit signals from the large
friction Tangential force that resists motion between two objects in class of seven-span receptors. The monomeric guanine-nucleo-
contact. tide-binding proteins that act in a wide range of intracellular
frond The leaf of a fern plant, which is often the most conspicuous signalling pathways are usually known as the ‘small GTPases’ or
part of the plant. ‘small G proteins’.
front focal plane The focal plane of a lens that lies in front of it G0 phase A phase of the eukaryotic cell cycle in which the cell
when viewed in the direction of illumination. This term is usually is temporarily or permanently arrested in its progression
employed when referring to the condenser lens, where the phase through the cycle and remains in a state resembling the G1
annulus is properly found. phase.

180
Glossary

G1 phase The ‘gap’ 1 phase of the eukaryotic cell cycle that gastritis Inflammation of the stomach.
succeeds cell division. It is the pre-DNA synthesis period of the gastrodermis See endoderm (2).
cell cycle in which the cell is increasing in size and chromosomes gastropods Snails, slugs, sea hares and kin (class Gastropoda,
are preparing for replication. phylum Mollusca), a class of molluscs typified (usually) by a
G2 phase The ‘gap’ 2 phase of the eukaryotic cell cycle that often twisted body and a helically coiled, nonchambered shell. A
succeeds DNA replication (S phase) and precedes mitosis (M characteristic feature is a large flat muscular ‘foot’ on which they
phase). crawl about. Present from the Cambrian to the present day.
GABA See g-Aminobutyric acid. gastrulation The period in animal embryonic development in
gain of function mutation A mutation in the protein-coding which tissue reorganization leads to the formation of the
region of a gene that confers an additional, deleterious, activity definitive germ layers (ectoderm, mesoderm and endoderm),
on the protein. Such mutations are generally dominant. formation of the gut, and the emergence of the body plan of the
galactomannan A polysaccharide composed of D-galactosyl and larval or adult organism.
D-mannosyl. In every case, the mannosyl units form the gating currents Transient currents caused by charge displace-
backbone structure (a linear main chain) with the D-galactosyl ments of voltage sensors within voltage-gated ion channels.
as single side units. gating The regulated opening and closing of the pore of an ion
gametangium (plural gametangia) A differentiated cell that channel.
produces gametes or whose protoplast functions in place of a gamete. gaussian Describes a generalized normal distribution, in reference
gamete The haploid germ cell produced by a sexually reproducing to the discoverer, Karl Friedrich Gauss (1777–1855).
organism. A gamete is capable of fusing with another gamete of
gaze Direction of the line of sight in space. Gaze depends on eye
opposite sex to produce a zygote. The male gamete is usually
position in the orbit and head position in space.
called the sperm, and the female the ovum.
G-banding Unique banding patterns that appear on chromosomes
gametic-phase disequilibrium An association (at greater than
stained with Giemsa dye.
chance levels) between particular alleles at two or more loci that
G-CSF See granulocyte colony-stimulating factor.
has arisen as a result of mating preferences, and thus as a result of
selection acting on the gamete. gel electrophoresis Procedure by which proteins or DNA or
gametogony Formation of gametes from haploid merozoites in RNA fragments can be separated according to size under the
parasitic protozoans. influence of an applied electric field. The matrix through which
gametophyte The phase of a plant’s life cycle in which the gametes the particles migrate is usually a polyacrylamide or agarose gel.
are formed. Male and female gametes fuse to form the sporophyte. gene Classically, the region of a chromosome that controls a single
c-Aminobutyric acid (GABA) One of two major inhibitory hereditary trait. The term gene is sometimes used as a synonym
neurotransmitters in the mammalian nervous system, the other for allele (a particular version of a gene) and sometimes as a
being glycine. synonym for locus (the position on the chromosome occupied by
cd T cells A subpopulation of T cells bearing a T-cell receptor the gene). At the molecular level, a gene is a sequence of DNA that
consisting of g and d chains. They are found in locations such as encodes the information for a protein or an RNA, together with
the epidermis. the regulatory sequences necessary for the gene’s expression.
gamone A soluble mating-inducing substance, also called mating gene amplification Sudden increase in the number of copies of a
pheromone. gene on a chromosome by successive rounds of gene duplication
gamont The cell (or organism) that produces gametes, or, in the that lead to a serial array of repeats of the gene.
case of ciliate conjugation, which produces gametic nuclei gene conversion (1) An unusual segregation of genetic markers in
(pronuclei). a single meiosis where three of the four gametes carry one allele
ganglion (plural ganglia) An anatomically discrete collection of and one carries the other, in contrast to the normal 2:2
nerve cell bodies within the central or peripheral nervous system, segregation. It is due to a nonreciprocal DNA recombination
containing from a few to many thousands of cell bodies and event that has converted one copy of one of the alleles into
associated structures. In invertebrates, a ganglion is usually a the other type. (2) A type of nonreciprocal recombination in
nodular mass surrounded by connective tissue. which a DNA sequence copied from a donor gene (or allele)
ganglion cells Neurons in the retina that carry visual signals to the displaces the equivalent sequence in a homologous acceptor gene
optic nerve. (or allele).
ganglioside A type of sphingosine-based glycolipid found espe- gene diversity The probability that two alleles chosen at random
cially in the cell membranes of nervous system cells. Production of from a population are not identical. It is also equivalent to the
gangliosides is also associated with certain carcinomas. proportion of heterozygous individuals in a randomly mating
gantry In computed tomography, the housing for the tube and population.
detectors. It contains all moving parts. There is a central aperture gene exchange Transfer of alleles between populations or species
through which the patient passes. by hybridization followed by backcrossing to the parental forms.
Garden of Eden hypothesis The hypothesis that modern humans This process is equivalent to gene flow but the term gene exchange
arose by a speciation event or something similar from a small focal is usually used to refer to interchange between genetically distinct
population. This is usually thought to have occurred in Africa. populations.
gas vesicles (1) Hollow, buoyancy-conferring, intracellular pro- gene expression The transcription and/or translation of a gene to
teinaceous structures found in a large number of aquatic and form the RNA and/or protein product. Regulation of gene
photosynthetic prokaryotes. (2) Intracellular vesicles produced expression can be at the level of transcription or translation.
by accumulation of a gas, e.g. hydrogen, which is produced as a gene flow Exchange of genes between populations caused by
byproduct of metabolic reactions. immigration and emigration of individuals across populations.

181
Glossary

gene-for-gene interaction In plant pathology, a specific recogni- genetic co-optation The stabilization, reinforcement or consoli-
tion between an avirulence gene from the pathogen and a dation by new or modified genetic circuitry of biological
corresponding resistance gene from the host plant, resulting in properties originally established by generic and epigenetic
resistance of the plant to that pathogen. mechanisms during organism–environment or tissue–tissue
gene-for-gene theory The principle that operates in some plant– interactions.
pathogen interactions, in which an avirulence gene in the genetic distance A measure of genetic or evolutionary closeness
pathogen encodes, or results in the production of, a molecule or remoteness between two taxa or sequences that originated
that is recognized by the product of a specific resistance gene in from a common ancestor. For DNA sequences it is the number of
the plant, which induces a resistance response in the plant. nucleotide differences per position between the two sequences.
gene fusion The covalent joining of DNA sequences from two genetic drift Random changes in allele frequencies in small
different genes, which may occur naturally or be constructed isolated populations as a result of factors other than natural
experimentally. selection, such as sampling of only small numbers of gametes in
gene gun A device used to accelerate and deliver DNA-coated each generation.
microprojectiles into a target tissue. genetic heterogeneity Similar phenotypes produced by defects in
gene knockout The result of creating a germline mutation in a different genes.
particular gene that completely disrupts the function of the genetic map Linear arrangement of genes and DNA markers
protein encoded by that gene. In mammals, the gene is disrupted on a chromosome, determined on the basis of recombination
by a technique that involves homologous recombination in frequencies.
embryonic stem cells in culture followed by preparation of genetic marker A detectable gene, DNA sequence or phenotypic
chimaeric mice by injection of these cells into a blastocyst. trait with a known chromosomal location.
gene pool (1) All the alleles present in a population of sexually genetic polymorphism The presence of two or more alleles of a
reproducing individuals. (2) All alleles at a given locus in such a gene at appreciable frequencies in a population.
population. genetic reassortment The generation of hybrid viruses possessing
gene rearrangement A site-specific somatic recombination event combinations of different genomic segments when a single cell is
that rearranges the order of genes or DNA sequences on the same infected with two or more related viruses containing segmented
chromosome. genomes, such as influenza viruses.
gene switching The process of turning a gene on or off as a result genetic screening The systematic search in a population for
of the binding of gene regulatory proteins to the regulatory individuals of particular genotypes.
elements of the gene. genetic testing Testing performed to determine the genotype of an
gene targeting See gene knockout. individual suspected of having a mutation.
genetically engineered live vaccine A vaccine containing a
gene therapy Treatment of genetically based disease by the
microorganism whose genes have been deliberately manipulated
introduction of a functional gene into the patient’s cells.
such that the infectious agent can still infect the host and induce
general circulation model A computer model capable of simulat-
immunity but cannot cause disease.
ing global climatological processes over long time scales.
genic Of or pertaining to individual genes.
general drift Changes in the genetic make-up of a population as a
genome (1) The total set of genes, and any additional non-coding
result of random processes.
DNA, carried by an individual organism, cell or virus particle.
general recombinase Enzyme that catalyses the homologous When the size of a genome is quoted, this usually refers to the
DNA pairing and strand-exchange reaction in general genetic haploid genome. (2) The total DNA content of a cell.
recombination. genome co-linearity The situation when homologous regions of
general-base catalysis Catalytic mechanism involving the chromosomes derived from two different species show the same
removal of a proton from a chemical group, which makes it order of markers or genes.
a stronger nucleophile. genome economization Reduction of genome size as observed in
generalist An organism with a broad realized niche, i.e. it is able to the genomes of animal mitochondria. Mitochondria are thought
live in a variety of different habitats or use a wide variety of to have reduced their genome sizes by transferring some of their
resources. genes to host cell genomes during the evolution of their symbiosis
generalized recombination See homologous recombination. with the host cell.
generative cells Cells within plasmodia that form sporoblasts and genome replication The stage of viral replication at which the
then spores. virus genome is copied to form new viral genomes.
genetic Pertaining to or caused by genes. genome sequence The complete nucleotide sequence of an
genetic balance The coordinated balance between the levels of organism’s DNA.
gene products in a particular genotype. Failure to establish genomic RNA RNA that is the genetic material of certain viruses
the correct balance usually results in severe defects and may be and encodes all their viral proteins.
lethal. genomics The study of a genome or genomes as a whole. The
genetic code The code by which the nucleotide sequence of a DNA genome-wide analysis of genes.
or RNA molecule specifies the amino-acid sequence of a genotoxic Able to cause damage to DNA.
polypeptide. It consists of three-nucleotide (triplet) codons that genotype The genetic make-up of an organism, as opposed to the
either specify a particular amino acid or a stop signal. phenotype, which is the outward appearance of the organism. A
genetic complexity The amount of genetic information carried listing of a complete genotype would include all alleles and
by an organism. For viruses lacking redundant nucleotide mutant sequences. In practice, genotypes are usually given in
sequences, complexity can be equated with genome size. respect of a selected set of loci under consideration.

182
Glossary

genotype–phenotype correlation The relationship of specific globular and fibrillar appearance At the electron microscopic
mutations at a genetic locus or loci to their manifestations in the resolution limit achievable with thin sections, concentrated
phenotype. solutions of small macromolecules are not resolved in detail
geochronological age The absolute age of a stratigraphical unit, but produce distinct ‘fibrillar’ or ‘globular’ aspects of such
horizon or body. It is usually determined by radioisotopic plasms.
methods and expressed in units of millions of years or millions of globular protein A protein with a relatively compact, roughly
years ago (Ma). spherical overall structure. Typically, hydrophobic amino acids
geognosy The study of the three-dimensional structure and are found in the center of globular proteins and hydrophilic
relationships of rock masses, particularly as applied to mines amino acids on the surface.
and mining. glucan A polysaccharide composed of glucose subunits.
geological record Rocks and fossils as found on the surface glucocerebroside A lipid that accumulates in Gaucher disease
of the Earth, from which the history of the Earth is inter- owing to a deficiency of the enzyme needed to degrade it.
preted. glucogenic Refers to metabolites that can be metabolized to
geological time The period from the formation of the Earth to the generate extra glucose or glycogen.
beginning of written history. glucogenic amino acid An amino acid that can be metabolized to
germ layers The three initial tissue types (ectoderm, mesoderm, give glucose.
endoderm) in an animal embryo, each of which gives rise to a glucomannan A polysaccharide composed of D-glucosyl and
particular set of organs and differentiated tissues. D-mannosyl units, generally with a preponderance of the
germinal centre An aggregation of rapidly proliferating activated latter.
B cells and a smaller number of CD4 helper T cells that forms in a gluconeogenesis The synthesis of glucose from molecules such as
lymphoid follicle of a secondary lymphoid organ (e.g. a lymph amino acids.
node) during an adaptive immune response. It is the site of B-cell glucose homeostasis The maintenance of plasma glucose con-
differentiation to plasma or memory cells and of isotype centration within normal limits (4.4–6.1 mmol l–1) by the actions
switching and affinity maturation of the humoral immune of regulatory mechanisms that compensate for changes in the
response. supply of and/or demand for glucose.
germination The outgrowth of a dormant reproductive propa- glucose–alanine cycle A metabolic pathway in which a product
gule, such as a spore or seed, into a new organism. from glycolysis (pyruvic acid) in muscle is converted to alanine
germline A cell lineage in animals that gives rise to the and transported to the liver, where it is converted to glucose,
reproductive (or germ) cells and is continuous through succeed- which returns to muscle.
ing generations. It separates early and clearly from somatic cell glutaminylglucan Polymer composed of glycosylated polygluta-
lineages during development. The cells of the germline are mine chains.
potentially immortal. glutathione (GSH) The tripeptide g-glutamylcysteinylglycine (g-
GFP See green fluorescent protein. GluCysGly), which is present at high concentration in cells. It has
ghost lineages, ghost taxa See range extensions. numerous biological functions in detoxification, reduction and
giardiasis Infection with Giardia lamblia, a pathogenic flagellate biosynthesis.
protozoan that can cause gastrointestinal symptoms in glutathione disulfide (GSSG) The symmetrical disulfide of
humans. glutathione.
Gibbs free energy A measure of energy that combines the glycan transferase An enzyme that transfers a specific sugar
enthalpy with the entropy of a system at constant pressure and residue from a nucleotide sugar onto other molecules such as
temperature. amino acids, lipids or sugars.
gingival hyperplasia Growth of the gingiva or tissue of the gums. glycocalyx The cell coat. In animal cells, a layer external to the
It is a frequent side-effect of therapy with dilantin or cyclo- plasma membrane that contains the carbohydrate moieties of
sporin A. glycoproteins and glycolipids embedded in the membrane, and
GISH See genomic in situ hybridization. secreted molecules such as proteoglycans.
glabella The upraised central part of the trilobite head. glycogen Polymer of glucose monomers linked by a-1,4-glucosidic
glacio-eustatic Global sea-level change caused by the growth or bonds and branched through a-1,6-glucosidic bonds. It is the
decay of ice caps. main glucose storage compound in animal cells.
glia Non-neuronal cells of the central nervous system, comprising glycogen granules Intracellular deposits of glycogen, a storage
microglia, astrocytes and oligodendrocytes. They provide var- polymer of glucose.
ious support functions required for the proper development and glycogenolysis The breakdown of glycogen into glucose.
function of neurons. glycolysis, glycolytic pathway The pathway by which glucose is
glioma Central nervous system tumour composed of and derived converted to pyruvic acid with production of energy in the form of
from glial cells. ATP, and reducing power in the form of NADH. It occurs in the
global alignment In DNA sequence analysis, the alignment of two cytosol and does not require oxygen. In aerobically respiring cells
or more sequences along their whole length. This form of it is the first stage in the complete oxidative breakdown of glucose.
alignment is not well suited to sequences of disparate length, but is In anaerobic cells, the term is often used for the complete pathway
well suited to the alignment of a group of sequence homologues. in which the pyruvate is subsequently converted to metabolites
global regulation Control of gene expression in a manner such as lactic acid.
appropriate to the prevailing nutritional and environmental glycopeptide resistance Resistance to one or more of the
conditions. glycopeptide antibiotics, e.g. vancomycin or teicoplanin.

183
Glossary

glycoprotein A protein to which short oligosaccharide side-chains gonopods Specialized limbs of myriapods, usually located
(chains of a few sugar residues) are covalently attached to the on or close to the reproductive segment (bearing the
protein backbone. The mass of sugar is normally a fairly small gonopore) and functioning in the handling of spermatophores
fraction of the mass of the whole molecule. and eggs.
glycosidase An enzyme that catalyses the hydrolysis of glycosidic goosecoid A homeobox-containing gene in Xenopus that encodes a
bonds, for example the bonds that link sugar molecules together transcription factor involved in executing the functions of the
in polysaccharides. Spemann organizer.
glycoside Molecules composed of a sugar linked to another GPI anchor See glycosylphosphatidylinositol anchor.
molecule (either another sugar or a nonsugar molecule) through GPI Glycosylphosphatidylinositol. See glycosylphosphatidylinosi-
a hydroxy group. A nonsugar component of a glycoside is called tol anchor.
an aglycone. G-protein-coupled receptors, G-protein-linked receptors A
glycosidic bond The covalent bond characteristic of sugar-to- large class of transmembrane receptor proteins in which binding
sugar linkages. of the extracellular ligand activates intracellular heterotrimeric
glycosome Cytoplasmic organelle in trypanosomatids in which proteins named G proteins. Activated G proteins stimulate
glycolysis takes place. intracellular enzymatic pathways or, in some cases, act directly on
glycosphingolipid Type of membrane lipid containing a ceramide nearby ion channels.
moiety linked through its terminal hydroxyl group to a variety of grade A group of animals similar in level of organization and
either monosaccharide or oligosaccharide units. adaptation. A level of evolutionary advance.
glycosylate To add a carbohydrate moiety to a protein biochemi- grading Formal scoring system for the cytological and architec-
cally. tural features of carcinomas.
glycosylated shell components Polypeptides with attached
graft-versus-host disease (GVHD) Disease that occurs when
carbohydrate moieties that surround carboxysomes.
mature immunocompetent T cells in a transplant react against the
glycosylation The covalent addition of oligosaccharide to another
tissues of the host. It is particularly a feature of bone marrow
molecule, usually a protein or lipid.
transplants into immunodeficient patients which are unable to
glycosylenzyme An enzyme on which a glycosyl group has been
reject the transplanted cells.
covalently bonded to a nucleophilic group.
Gram staining An important method for classifying bacteria
glycosylphosphatidylinositol anchor (GPI anchor) A covalent
which depends on the differential retention of Gram stain (crystal
linkage, by way of two fatty acids and an oligosaccharide chain,
violet) by the cell envelope. Bacteria that retain the stain are
between a membrane phospholipid (phospatidylinositol) and the
termed Gram-positive, and those that do not are termed Gram-
carboxyl terminus of an externally exposed protein.
negative.
glycosyltransferase An enzyme that catalyses the transfer of
Gram-negative Refers to bacteria that do not retain Gram stain
monosaccharide from an activated donor to a protein, lipid or
(crystal violet) (e.g. Escherichia, Pseudomonas).
oligosaccharide acceptor.
glyoxysomes A type of peroxisome found in plant cotyledons. Gram-positive Refers to bacteria that retain Gram stain (crystal
Although deficient in many peroxisomal functions, they do violet) (e.g. Clostridium, Streptococcus, Staphylococcus).
contain enzymes of the glyoxylate cycle. Gram-type A phylogenetic term used in the classification of
GlyR Abbreviation for the glycine receptor on neurons. prokaryotes, which is independent of the results of the Gram-
GMSF See granulocyte–macrophage colony-stimulating factor. staining reaction and is based on the presence or absence of
gnathobases Spiny processes at the bases of the paired legs of lipopolysaccharide in the cell envelope and on the 16S rDNA
arthropods, used in feeding. sequence analysis. Gram-type is defined as positive, negative or
gnathopods Mouthpart structures in some arthropods. zero (i.e. the archaea).
Golgi apparatus Organelle in eukaryotic cells that is responsible grana membranes Circular stacks of membrane formed by the
for polysaccharide synthesis, glycoprotein processing, lipid thylakoid membranes in chloroplasts.
biosynthesis and packaging of material for secretion to the granitic Describes rock composed of large crystals of
vacuole or cell surface. It receives proteins from the endoplasmic potassium feldspar, sodium plagioclase, feldspar, quartz
reticulum. The complete organelle consists of one or more stacks and biotite, formed from a magma rich in silicon,
of flattened sacs of membrane (cisternae). aluminium, potassium and sodium that cooled in a deep magma
Golgi body, Golgi stack Individual unit (stack of membrane sacs) chamber.
of the Golgi apparatus. Sometimes termed a dictyosome. granulation tissue The tissue that first forms during the repair of a
Golgi complex See Golgi apparatus wound or lesion. It consists of newly formed blood vessels,
gonad General name for organs in which gametes are formed activated fibroblasts and their products, and inflammatory cells.
(ovary and testis). granule An intracellular particle composed of material packed
Gondwana The southern continental landmass formed during the within a membrane vesicle.
early Mesozoic. It included what were to become South America, granule-mediated apoptosis Cell death that is induced by
Africa, Antarctica, Australia, India and New Zealand. perforin-mediated delivery of granzymes to the cytosol of the
Gondwanaland See Gondwana. target cell.
gonochoristic See gonochorous. granules of phagocytes Intracellular storage vesicles con-
gonochorous Describes organisms in which there is separation of taining microbicidal substances, present in phagocytic cells
the sexes such that individuals are either male or female (as such as macrophages and neutrophils. They can be mobilized to
compared to hermaphrode, where individuals produce both types deliver their contents to the exterior of the cell or into
of gametes). phagosomes.

184
Glossary

granulocyte (1) Type of white blood cell whose cytoplasm is rich in guard cells In plant cells, the paired cells comprising a stoma. Their
granules. In humans, there are three main types – eosinophils, expansion as a result of a change in turgor pressure opens the
basophils, and neutrophils (polymorphonuclear leucocytes) – the stomatal pore.
most abundant of which is the neutrophil. (2) Major haemocyte guide fossils A fossil species that characterizes a particular interval
type found in all major arthropod taxa and characterized by the of the rock record, which is termed the biozone of that species.
presence of numerous prominent intracytoplasmic membrane- They are essential tools in local and international biostratigraphic
bounded granules. correlation.
granulocyte colony-stimulating factor (G-CSF) A cytokine that Guillain–Barré syndrome Acute inflammation of peripheral
predominantly stimulates the growth of granulocytes from bone nerves following an infectious disease of viral or bacterial
marrow cells. origin.
granulocyte–macrophage colony-stimulating factor Gulf War syndrome A group of ailments reported by a number of
(GMSF) A haematopoietic growth factor that stimulates growth soldiers who fought in the war to liberate Kuwait from its Iraqi
of granulocytes (G) and macrophages (M) from bone marrow invaders. The cause remains unknown and there is controversy
cells. over the syndrome’s existence.
granulomatous disorders Diseases characterized by the appear- gumma The tertiary noninfectious syphilis lesion that occurs
ance of granulomas. either at the site of inoculation in the sensitized host or
granzymes A family of cytotoxic serine proteases that are present following spread of Treponema pallidum through the blood
in the granules of cytotoxic cells. stream.
Graves disease An autoimmune disease caused by the production gustatory Relating to the sense of taste.
of antibodies that stimulate the thyroid-stimulating hormone gustducin The a subunit of a G protein that is found uniquely
receptor, resulting in an overproduction of thyroid hormones in a subset of taste receptor cells and in certain chemoreceptor
(thyrotoxicosis). cells in the gut. Gustducin is hypothesized to couple bitter
receptors to intracellular signalling pathways inside taste receptor
green fluorescent protein (GFP) An autofluorescent protein
cells.
from the jellyfish Aequorea victoria. It is commonly used, via
guttation The excretion of water from specialized openings, the
recombinant DNA techniques, to mark a particular protein
hydathodes, in plant leaves when the air is saturated.
within living cells and/or whole organisms, so that it can be
detected visually. Gvp proteins Proteins associated with gas vesicle formation.
gymnamoebae A group of amoebae that lack a definite enclosing
greenstone Basalt that has cooled under the ocean, usually
shell or test, but sometimes have a thin glycocalyx coating the
forming lava pillows, and then is later subjected to low-grade
plasma membrane.
(greenschist-grade) metamorphism, forming the green mica
gymnosperm A plant that produces seeds not enclosed in an
chlorite that gives the characteristic colour.
ovary. Gymnosperms include the conifers.
grey matter A generic term for a collection of nerve cell bodies in
gynoecium All of the female reproductive structures in a
the central nervous system.
flower.
grey-scale image An image in which the intensity of signal coming
from, or the amount of sound or radiation absorbed by, each
region of the body is represented on an image of the body by a
shade of grey ranging from black to white.
H See enthalpy.
group selection Differential survival or reproduction of groups of
organisms, such as a population of parasites on a host. HAART See highly active antiretroviral therapy.
habitat Both the place occupied by an organism and the combina-
growth cone A specialized region at the tip of a growing neurite
tion of physical and environmental conditions required for an
(axon or dendrite) that is responsible for sensing the local
organism to survive and reproduce.
environment and moving toward the neuron’s target cell. Growth
habituation A reduction in response following repeated presenta-
cones are roughly hand shaped, and extend several actin-based
tions of a stimulus.
processes, called filopodia, that navigate the axon tip over the
substrate. haem The iron(II) complex of a porphyrin, a prosthetic group on
many proteins
growth factor receptors Transmembrane proteins in the plasma
haemagglutination The clumping of red blood cells, usually by
membrane that bind growth factors and mediate their biological
specific antibody.
effects.
haemagglutinin A protein that can bring about the agglutination
growth factors General term for cytokines that affect cell
of red blood cells. Influenza haemagglutinin is a glycoprotein on
proliferation and differentiation.
the surface of the influenza virus that is responsible for binding of
Gs G proteins that activate adenylyl cyclase. the virus to receptors on host cells.
GSH See glutathione. haematite An iron oxide mineral, typically red in colour;
GSK-3 Glycogen synthase kinase 3, a serine/threonine kinase with a Fe2O3.
role in early embryonic development. It is regulated by multiple haematocrit The proportion of red cells in the circulating
signalling pathways, including Wnt signalling and insulin blood related to the plasma volume, often reported as the
signalling. percentage of a centrifuged blood sample that contains the red
GSSG See glutathione disulfide. cells.
guanine Commonly occurring purine base, one of the four types of haematogenous dissemination Spread of a pathogen via the
bases in RNA and DNA. blood system.

185
Glossary

haematopoiesis The production of all the cellular elements of haploid An individual or cell (e.g. a gamete) having a single set of
blood, including platelets, red blood cells and leucocytes. chromosomes (n). The number of chromosomes is species
haematopoietic cell A cell from which blood cells derive. specific.
haemin The chloroiron(III) complex of a porphyrin. haploinsufficiency Insufficiency of gene product resulting from
haemocoel (1) A blood-filled body cavity that bathes the internal the presence of only one functional copy of the gene instead of the
organs. (2) The open blood system of arthropods and molluscs. It usual two.
comprises a system of sinuses through which the haemolymph or haplorrhine A cladistic term of primate classification that includes
‘blood’ travels. tarsiers and anthropoids, in order to better reflect their shared
haemoglobin A haem-containing protein, contained in red blood phylogenetic origin.
cells in vertebrates, that reversibly binds molecular oxygen and haplotype The combination of the major histocompatibility
transports it from the lungs to tissues. complex (MHC) alleles carried on one chromosome of the
haemolymph The multipurpose ‘blood’ of arthropods. It trans- homologous pair.
ports cells, nutrients, wastes and dissolved gases throughout the hapten A compound, usually of low molecular weight, that is not
body. itself immunogenic but that, after conjugation to a carrier protein
haemolysin A protein that causes the rupture of red blood cells. or cell, becomes immunogenic and can induce specific antibodies.
haemolysis The lysis of red blood cells. These antibodies can bind the hapten alone in the absence of the
haemolytic Causing lysis of red blood cells. Many pathogenic carrier.
bacteria, notably Streptococcus pyogenes and some other
haptocyst Organelle in the suctorian tentacle that is extruded to
streptococci, possess this property.
make contact with a prey organism.
haemolytic anaemia Anaemia due to decreased survival of red
haptotaxis The directional migration of cells towards the source of
blood cells in the circulation. Clinically, the disease is character-
a chemical signal bound to a substrate.
ized by anaemia with a raised reticulocyte count in the absence of
bleeding. hard selection Selection based on the absolute fitness of indivi-
haemolytic disease of the newborn More correctly, haemolytic duals in a population. It is usually independent of the frequency of
disease of the fetus and newborn. A fetal and neonatal anaemia particular genotypes.
caused by the action of maternal antibodies against fetal red hardening Acclimation of a chill-resistant plant to freezing
blood cells. temperatures by pretreatment at low, but not freezing, tempera-
haemophilia A hereditary bleeding disorder that affects almost tures.
exclusively male patients. It is caused by genetic defects that result haustorium (plural haustoria) A specialized branch of a fungus
in the absence of either of the clotting factors VIII or IX from the developed within the cell of another fungus or plant, which
plasma. absorbs nutrients.
haemorrhagic fever A disease characterized by widespread H1 ATPase A transmembrane protein that uses the energy released
bleeding as a result of blocking and permeabilization of by hydrolysis of ATP to transport H1 across a membrane.
1
capillaries throughout the body. H -dependent vesicular uptake Uptake of neurotransmitter into
haemostasis The physiological mechanisms that normally main- synaptic vesicles by transporter proteins that use the proton
tain the blood in a fluid state but immediately arrest of bleeding gradient between the cytosol and the lumen of the synaptic
from a damaged vessel by blood clotting, vasoconstriction or vesicles as an energy source.
other means. H1-pyrophosphatase (H1-PPase) Pyrophosphate-driven proton
hairy root Aberrant root growth caused by the bacterium pump localized in the tonoplast of plant cells.
Agrobacterium rhizogenes on dicotyledonous plants. HCG See human chorionic gonadotrophin.
half-centre oscillator A reciprocally inhibitorily coupled pair of headful packaging A mechanism of encapsulation of DNA in a
circuits that oscillate in anti-phase. phage head in which a concatemeric DNA is cut after uptake of a
half-life The time taken to reduce the amount or activity of a given length of DNA sufficient to fill the head.
substance by half. Used, for example, as an estimate of the time a
headgut Mouthparts and pharynx of the mammalian intestinal
drug remains in the body, or as a measure of the radioactivity of a
system.
substance. In transplantation medicine it refers to the time at
heart A valved, muscular pump in animals that propels fluid,
which half of the patients have lost their grafts.
usually blood or lymph, around the body.
halides The negatively charged monovalent anions F, Cl, Br,
I . heat-shock protein Protein synthesized in response to raised
hallucinogen A substance that causes mental disorientation temperature or other stress. Heat-shock proteins assist the correct
(hallucinations). folding of other proteins newly synthesized and denatured by
halocarbon A (usually manmade) organic compound containing stresses.
halogen atoms, e.g. pentachlorophenol, a wood preservative. heavy chain The larger of the two types of polypeptide chain in an
They are often highly resistant to biodegradation. immunoglobulin molecule.
halophile An organism that requires high concentrations of salt to Hebb rule Synapses should be strengthened if there is both
survive. presynaptic activity and strong enough depolarization of the
halotolerant Capable of growth in hypersaline conditions but not postsynaptic cell to cause spiking.
necessarily requiring them for growth. helicase An enzyme that unwinds double-stranded DNA in an
hammerhead structure A particular class of ribozyme present in ATP-dependent reaction.
one or both polarity strands of some viroid and viroid-like helix-turn-helix A common protein fold associated with the ability
satellite RNAs. to bind DNA.

186
Glossary

HELLP syndrome Complication of pre-eclampsia characterized by heteroduplex (1) Double-stranded DNA in which the two strands
haemolysis, liver damage and disseminated intravascular coagu- are not of identical complementary sequence. (2) A double-
lation. stranded nucleotide polymer in which one strand is DNA and one
helper component A nonstructural protein encoded by the virus strand is RNA.
genome that is needed for a successful transmission of the virus by heteroencapsidation The situation when some of the RNA of one
the insect. virus (or virus strain) is coated with proteins of another virus (or
helper T cells A functional subset of T cells which carry the CD4 strain).
co-receptor and primarily function to help B cells produce heterogametic Having different kinds of sex chromosomes, e.g. X
antibody. and Y in mammals. In mammals, males are the heterogametic sex.
hemizygous Describes the condition in which only a single copy of heterogenesis See alternation of generations.
a set of genes is present in a diploid organism. Human males are heterograft See xenograft.
hemizygous for the genes on the X chromosome because they heterokaryon A multinucleate cell or mycelium having genetically
possess only one X. different nuclei.
heparin Sulfated glycosaminoglycan of varying molecular weight heterologous Pertaining to a gene or protein which is being
which is composed of polymers of repeating disaccharide units of expressed in an organism or cell in which it is not normally
glucosamine and D-glucuronic acid or L-iduronic acid. It is present.
released by mast cells and basophils and has anticoagulant heterologous expression Expression of a foreign gene in a given
activity. expression system. An example is when a human gene is expressed
hepatitis Any disease characterized by inflammation and damage in Escherichia coli.
to the liver. heteromeric A protein composed of two or more different
hepatocyte The main cell type in liver, which carries out most of polypeptide chains.
the liver functions. heteroplasmy The presence of two or more subpopulations of
hepatocyte growth factor (HGF) Secreted signalling molecule mitochondrial DNA in the same organism or cell.
expressed in the vertebrate embryo in the lateral plate mesen-
heteropolysaccharide A polysaccharide composed of more than
chyme and required for the migration of prospective muscle cells
one type of monosaccharide.
into the limbs.
heterosaccharide A polysaccharide composed of a complex
hepatosplenomegaly Enlargement of the liver and spleen
mixture of monosaccharides.
due to blood production in these organs and congestive cardiac
heterosis The case when the heterozygous progeny is superior in
failure.
fitness to its homozygous parents.
hepatotoxicity The property of being toxic to liver cells.
heterospecific Between different species.
herbivore An organism that consumes live plant tissues.
hermaphrodite An animal that produces both male and female heterosynaptic long-term depression (LTD) The weakening of
inactive synapses as a result of activity in other synapses on the
gametes. Self-fertile hermaphrodites can produce viable embryos
by fertilizing their own eggs with their own sperm. same cell.
hermaphroditism Being male and female at some stage in the life heterotaxia, heterotaxy (1) Randomized left–right distribution
cycle. Often both male and female organs can be present at once, of organs. (2) A cardiac looping defect characterized by altered
and this is termed simultaneous hermaphroditism. left–right asymmetry that can be expressed as randomization
herpes simplex virus A herpesvirus producing a variety of (situs ambiguus) or complete reversal (situs inversus) of normal
infections involving mucocutaneous surfaces, the central nervous organ positions (situs solitus).
system and visceral organs. heterothallic Describes organisms in which matings are restricted
herpes An infectious disease caused by a herpesvirus. to parents of different mating types.
herpesviruses Viruses of the family Herpesviridae which typically heterotrichous Having two (or more) forms of filament develop-
cause cold sores and genital sores. ment in the same thallus.
hesperornithiforms Flightless, toothed birds from the Cretaceous heterotrimeric G proteins G proteins that consist of three
of North America and Europe, including Hesperornis regalis, different subunits (a, b and g) and are activated by receptors of
Parahesperornis alexi, and Baptornis advenus. the seven-span class. Four major classes of G proteins are Gs, Gi,
heterochromatin Densely staining chromatin that remains Gq and Go, which interact with distinct downstream effectors
relatively condensed throughout interphase, and is later (enzymes and ion channels).
replicating than euchromatin. It is found at centromeres and heterotroph Organism that requires organic carbon (usually
telomeres, is usually gene poor, and is thought to be genetically material derived from other organisms) for energy and growth.
inactive. heterotrophic Requiring organic compounds as a source of
heterochronic Describes evolutionary change that occurs by carbon.
modifying the timing of developmental events. heterotropic interactions Allosteric interactions between sites
heterochrony The slowing-down or speeding up of the relative that bind different types of ligands (e.g. between O2 and H1 or
rate of development of an organ (or whole organism) by 2,3-BPG in haemoglobin).
comparison with its development in the ancestor. It is an heterotypic pairs Conjugating cells of different mating type.
important process in evolution. heterozygosity A measure of genetic variation within a popula-
heterocyst A terminally differentiated nitrogen-fixing cell present tion. For a given locus, the frequency of heterozygotes at that
in many cyanobacteria. locus within the population.
heterodimer A protein molecule composed of two different heterozygote A diploid organism that carries two different alleles
subunits. at a given genetic locus.

187
Glossary

heterozygote advantage Situation in which a heterozygote has histopathology The study of the structural alterations of cells and
greater fitness than either of the homozygotes in a given tissues caused by disease.
environment. Also known as hybrid vigour and heterosis. HLA Human leucocyte antigen, the name given to the human major
heterozygous Possessing two different alleles at a given genetic histocompatibility complex.
locus. hnRNA Heterogeneous nuclear RNA, an old name for pre-mRNA.
heuristic search In taxonomy, choosing a phylogenetic tree based hnRNP protein Heterogeneous nuclear RNA protein. A protein
on the examination of only a selected subset of all possible trees. that binds pre-mRNA in the nucleus. Although a distinction is
The tree chosen may not be the optimal tree. sometimes made between hnRNP proteins and splicing factors,
HEV See high endothelial venule. there is considerable overlap due to incomplete knowledge and
hexagonal Describes a form of crystal symmetry characterized by actual overlap in function.
three equal axes intersecting at 1201 and a fourth of unequal H-NS A bacterial histone-like protein.
length perpendicular to the others. This type of symmetry is hodological Refers to axonal pathways and connections in the
typical of quartz and calcite. central nervous system. Derived from the Greek hodos, which
Hfr A bacterium with a sex plasmid such as F incorporated into means way or road.
its chromosome and which transfers its chromosome at high HO-endonuclease A restriction enzyme from the yeast Saccharo-
frequency. myces cerevisiae that recognizes a single site in the yeast
HGF See hepatocyte growth factor. genome.
hiatus A gap within the time record represented by a stratigraphical Holliday junction A four-way junction linking two DNA mole-
succession. cules which results from the reciprocal exchange of DNA strands
hibernaculum Small resting body of entoprocts (Entoprocta) between homologous DNA duplexes. It is a central intermediate
consisting of an inner mass of cells, usually with much yolk, in homologous recombination. Resolution of the Holliday
surrounded by a thick chitinous wall. junction by specialized nucleases can give rise to crossover
hierarchy A system with levels, such as the familiar Linnaean recombinants.
hierarchy in biological classification. holobasidium A single-celled basidium, often club shaped.
high endothelial venule (HEV) A specialized blood vessel holocentric chromosome A chromosome that has a single and
through which lymphocytes leave the blood circulation and enter continuous (diffuse) centromere, extending along the whole, or a
lymphoid tissue. considerable part, of the chromatids and to which the spindle
higher-order repeats Tandem arrays in which basic repeating microtubules attach.
units are combined into composite units such as dimers. holoenzyme For multisubunit enzymes, the complete enzyme
highly active antiretroviral therapy (HAART) Therapy usually containing all the components necessary for its regulated
involving a combination of at least three antiretroviral drugs, activity.
typically two reverse transcriptase inhibitors and a protease hologenous The situation where the inner and outer walls of
inhibitor. the conidiogenous cells and the developing conidium are
high-performance liquid chromatography (HPLC) Chromato- continuous.
graphy performed under high back pressure in steel columns HOM genes The name given to the Hox genes in Drosophila.
using a matrix of silica particles of different sizes that enables home range Area used over extended periods by an individual or
separation of polypeptides according to their size, net charge or social group for feeding and other needs.
hydrophobicity. homeologous Refers to nucleic acid sequences that are similar but
high-threshold channel A voltage-gated channel that opens not identical.
when the membrane potential is depolarized close to 0 mV. homeostasis The intrinsic physiological process by which the
hilum A small gap or hollow in an organ where vessels, nerves and internal systems of the body (e.g. ionic and osmotic concentra-
ducts enter or leave it. tions, hydration, pH, resting body temperature) are maintained
hindbrain The posterior part of the brain, consisting of the pons, at equilibrium (autoregulated), despite variations in the external
medulla and cerebellum. environment.
hindgut The large intestine in many species. homeothermic ‘Warm-blooded’, able to maintain a relatively
hirsutism Presence of excessive body and facial hair. constant body temperature under a range of environmental
HIS See Hospital Information System. temperatures.
histidine protein kinase Enzyme with a conserved histidine homeothermy An old term used to refer to animals that are able to
residue which, when activated, can autophosphorylate at the maintain stable, high body temperatures. This word has generally
histidine and then phosphorylate target proteins. been replaced by the terms ‘endothermy’ and ‘exothermy’, which
histiocyte Tissue cell derived from the monocyte which can be emphasize the functional nature of the regulation rather than its
activated to become a phagocyte. result.
histoincompatibility The genetic difference between individuals homeotic (1) Describes genes that specify the identity of parts of
that is responsible for the rejection of skin and organ grafts. the body. (2) Describes mutations in such genes that transform
histolysis Tissue breakdown due to remodelling of extracellular one body part into another.
matrix by proteases. homeotic transformation The replacement of an organ with an
histones Small basic proteins that are a major component of organ of different type during development.
chromatin. They form the cores of nucleosomes, in which DNA is homing Targeted migration of lymphocytes to a given anatomical
wrapped around the protein core. location.
histone-like proteins Bacterial proteins that can bend or package homing receptor An adhesion molecule on lymphocytes that is
DNA. required for lymphocyte migration to a specific target organ.

188
Glossary

Homininae, hominins The name given to a tribe of higher homotetramer A tetrameric molecule composed of four identical
primates that includes humans. There is only one living, subunits.
monotypic genus, Homo sapiens, but the fossil record of the homothallic Not sexually differentiated, applied to populations of
group includes species attributed to Australopithecus, Paranthro- e.g. yeasts of the same mating type.
pus and Ardipithecus. homotropic interactions Allosteric interactions between sites
Hominoidea Superfamily of primates that includes all living and that bind the same type of ligand (e.g. O2 in haemoglobin).
fossil apes. homotypic pairs Conjugating cells of the same mating type.
hominoid A general term of primate classification that refers to homozygote A diploid organism that carries two identical alleles
apes and humans. at a given genetic locus.
homoctenids The Cricoconarida, tiny, cone-shaped fossil organ- homozygous Possessing two identical alleles at a given genetic locus.
isms whose mode of life was similar to that of modern planktonic horizontal gene transfer (1) The transmission of genetic material
gastropods such as pteropods. between cells in a process not involving reproduction. (2)
homodimer A protein composed of two identical subunits. Transfer of genetic material between species.
homogalacturonan A polysaccharide composed of 1,4-linked horizontal transport Concentration or dispersal of surface algae
galactosyluronic acid residues. by small- to large-scale currents.
homogametic sex The sex with two of the same kind of sex host (1) The symbiotic partner that is the largest or most
chromosome (e.g., mammalian females, which are XX). conspicuous member of the association. (2) Animal or plant
homogeneity Consisting of identical entities. A substance or cell infected by a virus, bacterium, fungus or other parasite.
culture is homogeneous when exacting analytical tests show that host factors In insect immunity, factors secreted by the immuno-
it is pure. cyte, that opsonize the foreign antigen and attract other
homogeneous RNA RNA of a single length and sequence. immunocytes.
homograft See allograft. host switch Transfer of a parasite from one host to another.
homoiohydric Describes organisms that are resistant to desicca- Hox genes Family of homeobox-containing genes that are
tion and maintain internal hydration, even in drought, by the use characteristic of animals and are involved in specifying positional
of waterproof cuticles, etc. identity along the anterior-posterior axis.
homologous (1) Refers to DNA sequences, molecules or
HPLC See high-performance liquid chromatography.
structures that are similar as a result of their derivation
HTLV-I Human T-lymphotropic retrovirus type-I (HTLV-I). It
from a common ancestor. (2) Refers to the maternal and
causes leukaemia and lymphomas.
paternal copies of a given chromosome in a diploid organism,
HU A bacterial histone-like protein.
which carry the same genetic loci although they may carry
human chorionic gonadotrophin (HCG) Polypeptide hormone
different alleles.
that maintains the corpus luteum in humans.
homologous DNA pairing and strand exchange The enzymatic
human genome All the genes possessed by a human individual.
process that pairs an initiating single-strand DNA substrate with
a homologous duplex during homologous recombination. Human Genome Project International project to map and
homologous recombination Reciprocal exchange of genetic sequence the human genome.
information between two homologous chromosomes, or between humification The biochemical reprocessing by microbes of or-
DNA molecules sharing extensive homology. ganic matter in leaf litter, particularly the more resistant tannins
homologous sequences DNA or amino-acid sequences with a and lignins, into complex aromatic compounds, collectively
common origin. They result either from speciation or gene termed humic acids or humus.
duplication. humoral (1) Relating to any clear fluid or semifluid translucent
homology (1) Relationship between characters or character states anatomical substance. (2) Relating to the B-cell or antibody
in organisms that derive from the same feature in a common response of adaptive immunity.
ancestor. (2) Significant sequence identity between genes or humoral capsule See encapsulation.
proteins at the nucleotide or amino acid level, respectively, as a humoral factors All substances (e.g. attacins, cecropins, defensin,
result of their derivation from the same ancestral sequence. gallysin 1 and 2, haemolin and lysozyme) that are synthesized by
homology-dependent gene silencing A type of epigenetic gene the insect’s immune system and the fat body in arthropods in
silencing induced by the presence of several homologous copies of reaction to invasion by foreign antigens.
a gene. humoral immunity Antibody-mediated immunity.
homomeric Describes a protein composed of two or more humoral responses The immune response mediated by B cells.
polypeptide chains of identical sequence. Huntington disease A heritable degenerative brain disease.
homonomous Having segments that are very much alike. HVRI One of the regions within the mitochondrial DNA that codes
homoplasmy The case where all mitochondrial DNA molecules in for proteins involved in cellular metabolism.
a population are identical. hybrid (1) The offspring of a cross between two pure-breeding lines
homoploid Possessing the same number of sets of chromosomes. of different genotype. (2) The offspring of a cross between
homopolymers Nucleotide polymers composed of only one type different species.
of nucleotide. hybrid zone A region where the geographic ranges of two
homopolysaccharide Polysaccharide consisting of one type of genetically divergent groups of populations meet and produce
monosaccharide subunit. at least some offspring of mixed ancestry.
homosynaptic long-term depression (LTD) The situation when hybridization (1) A cross between individuals from genetically
the long-term decrease (depression) in synaptic strength occurs at differentiated populations. (2) The pairing of two complementary
the same synapses that were stimulated. nucleic acid strands from different sources.

189
Glossary

hybridization probing The use of a labelled nucleic acid molecule hyperacute rejection A rejection response against a transplanted
as a probe to identify complementary sequences to which the organ that is initiated by pre-existing host antibodies to antigens
probe base pairs. on the transplant. Rejection occurs rapidly, within minutes to
hydration shell Water around a solute that is perturbed by the solute. hours after transplantation, and is untreatable.
hydration Solvation in water. hyperammonaemia Increased blood ammonia level that may
hydrocephalus Accumulation of intracranial cerebrospinal fluid have a potentially toxic effect on the brain.
(CSF) resulting in ventricular enlargement, and, frequently, hyperbilirubinaemia Increased bilirubin concentration in the
increased intracranial pressure. circulation due to haemolysis (breakdown of red blood cells),
hydrodynamic Related to the force or pressure of moving water releasing haem from haemoglobin, which is broken down to
(or other fluids). bilirubin.
hydrodynamics The motion of fluids. In sedimentation this refers hypercalcaemia A condition in which there are relatively large
to the motion of particles dissolved or suspended in a liquid. amounts of calcium ions in body fluids.
hydrogen bond Noncovalent interaction involving two electro- hyperchromic shift, hyperchromicity Increase in absorption of
negative atoms (in biochemistry mainly N, O and S), one acting as ultraviolet light of 260 nm wavelength that occurs when a DNA
a donor and the other as an acceptor of a hydrogen atom. An duplex dissociates.
important type of bond in biological macromolecules. hyperglycaemia Increased level of blood sugar.
hydrogenosome A membrane-bounded respiratory organelle, hyperlipidaemia Increased levels of blood lipids.
typically found in anaerobic protozoa, in which enzymes oxidize hypermodified base Highly modified base in a nucleic acid.
pyruvate to acetate and carbon dioxide. Under anaerobic hyperosmolality Having a greater osmotic pressure than body
conditions this reaction releases hydrogen. fluids.
hydrolases Enzymes that cleave ester, amide, anhydride and hyperphenylalaninaemia An error in metabolism whereby
glycosidic bonds by using the oxygen of water as the nucleophile dietary phenylalanine is not converted to tyrosine and so its
in a substitution reaction. concentrations build up to dangerous levels in the blood.
hyperplasia Increase in the size of a tissue or organ resulting from
hydrophilic Easily dissolvable in and attracted to water, as a result
an increase in the number of cells.
of polar or ionic interactions. Hydrophilic chemical structures
hyperpolarization A change in membrane potential away from
include hydrogen-bonding groups and charged groups, which
zero, making it more negative.
can participate in the structure of water.
hypersaline Having a salinity greater than that of sea water.
hydrophilicity The property of having high solubility in water.
hypersensitive response (1) A manifestation of plant disease
hydrophobic Water-repelling. Nonpolar hydrophobic molecules or
resistance resulting from restriction of pathogen to a small zone
chemical groups tend to be oriented toward the interior of proteins
around the infection site and accompanied by localized death and
or toward the interior of a lipid bilayer membrane, and are insoluble
necrosis of host tissue. (2) An exaggerated inflammatory response
in water but soluble in lipids. Hydrophobic chemical structures,
to an antigen.
such as chains of methylene and methyl groups, cannot participate
hypersensitivity An exaggerated inflammatory response to the
in the structure of water, and thus force an energetically
exposure of an antigen.
unfavourable organization upon the water structure.
hypertension Increase in blood pressure.
hydrophobicity Low solubility in water. hyperthermophile A microorganism with a temperature opti-
hydrophobicity plot Diagram showing the running average of the mum for growth above 801C.
index of hydrophobicity of amino acids in a protein sequence. hypertrophy Increase in the size of a tissue or organ resulting from
Hydrophobic stretches of amino acids tend to arrange in such a an increase in the size of the cells present.
manner as to avoid contact with polar charges. hyperuricaemia Increased concentration of uric acid in peripheral
hydrops fetalis A condition that arises in the fetus as a result of blood plasma.
extreme anaemia. It results in lack of oxygen in the fetal tissues hypervariable regions The parts of an immunoglobulin variable
and congestive cardiac failure, leading to a bloated appearance region having the highest degree of amino-acid sequence
with swelling of the tissues. polymorphism and principally responsible for conferring speci-
hydrostatic equation dF/dp 5 – a, where F is the geopotential of ficity on the antigen-binding site.
an isobaric surface, p is pressure, and a is specific volume. hypha (plural hyphae) The fungal vegetative growth form, which
hydrostatic skeleton A fluid-filled cavity in e.g. echinoderms is a tubular branching filament that elongates by apical extension.
which is constrained by the muscular body walls. hyphal body A segment of a hypha that becomes separated before
hydrothermal system An ocean subsurface at which magma- reproducing by budding or fission.
heated water spews out at temperatures as high as 3501C. hyphosphere The zone immediately surrounding the external
hydrothermal vents Hot springs found at depths up to 4000 m in surface of the hypha.
areas along mid-ocean ridges in the Atlantic and Pacific Oceans hypoblast See primitive endoderm.
and support a unique community of organisms. They are created hypodermis Epidermis or skin of a nematode.
by plate movements that occur on the crustal zones of the Earth. hypodigm All the fossil specimens that have been attributed to an
hydroxyapatite A naturally occurring calcium phosphate extinct species.
(Ca10(PO4)6(OH)2), which is the major inorganic constituent hypoglossal nucleus Region of the medulla that controls the
of the bone matrix. muscles of the tongue and syrinx (vocal organ of birds).
hyoid apparatus Bones in the neck of tetrapod animals that hypoglossal Nerve that supplies the intrinsic muscles, and all but
support the tongue and larynx. They are the remains of one of the extrinsic muscles, of the tongue.
embryonic gill arches. hypokalaemia Low level of potassium in the blood.

190
Glossary

hypoplasia Underdevelopment of a tissue. IL-5 See interleukin-5.


hypoproteinaemia Low plasma protein levels in the blood. IL-6 See interleukin-6.
hyposplenism Reduction in activity of spleen. ileus Obstruction of the bowel.
hypostome Most apical region in hydra, surrounding the mouth image artefact A component or aspect of an image that does not
opening. correspond to a real object or function.
hypothalamus An area of the brain situated below the thalamus imaginal disc A single-layered pouch of undifferentiated cells in
containing centres regulating many functions, including repro- insect larval stages that gives rise to adult structures such as the
duction. wings, legs, antennae and eyes.
hypotonic Refers to a fluid whose osmolarity is lower than the imidazole An azole antifungal agent containing an azole ring with
osmolarity of normal extracellular fluid. two nitrogen atoms. An example is ketoconazole.
hypoxaemia Decreased oxygen tension (concentration) in the
imino group A portion of a molecule with the formula 5C5NH.
blood, measured by arterial oxygen partial pressure values. It is
immediate early gene A member of a set of genes that is rapidly
sometimes associated with decreased oxygen content.
but transiently expressed in response to extracellular signals.
hypoxia Condition in which the concentration of oxygen is greater
immediate hypersensitivity Immunological hypersensitivity
than 0% but less than 20%. In water this is a concentration of less
reaction that occurs in a sensitized individual within minutes
than 2 mg O2 per litre.
to a few hours of re-exposure to antigen. Most immediate
hypersensitivity reactions are mediated by IgE.
immune activation Enhancement of the antimicrobial activities of
in situ In the natural location.
macrophages by interferon-g produced by antigen-activated T cells.
in vivo gene transfer Gene transfer directly into cells of the body.
immune evasion Strategy by which microbial pathogens subvert
iatrogenic Induced by medical intervention or treatment.
host immune responses in order to establish an infection, e.g. by
IC50 Inhibitory concentration 50. The drug concentration that
expression of virulence factors such as capsules, toxins, adhesins,
inhibits viral activity by 50%.
siderophores, modulins, impedins.
iccosomes Immune complex-coated bodies released by follicular
immune haemolytic anaemia Anaemia due to destruction of red
dendritic cells in germinal centres. Iccosomes are endocytosed by
blood cells mediated by an immune reaction, i.e. by antibody,
antigen-specific B cells, processed and presented to T cells in the
with or without the involvement of complement.
germinal centre.
ichthyornithiforms Toothed flying birds from the Late Cretac- immune repertoire The totality of lymphocyte receptors. The
eous of North America, including Ichthyornis dispar and repertoire consists of millions of precursors of mature lympho-
Ichthyornis victor. cytes, each bearing antigen receptors of a unique specificity.
ICM See inner cell mass. immunity Protection against a particular disease. Innate immunity
icosahedral Having 20 sides or faces. is that nonspecific protection conferred by the body’s fixed
icosahedron A regular polyhedron with 20 faces, each an defences such as the skin, antimicrobial proteins and phagocytes.
equilateral triangle, and 12 vertices, found as the morphological Long-lasting adaptive immunity is generated by the immune
form of many viruses. system after a first exposure to the disease, and is specific for the
pathogen or toxin.
icterus gravis neonatorum Yellow jaundice in the newborn due
to haemolysis. immunization Introduction of an immunogen into a host in order
identity elements A set of spatially organized chemical features of to initiate an immune response against the immunogen.
an RNA molecule specifically recognized by an enzyme or immunoadsorbent A material carrying bacterial proteins A or G
protein. (from Staphylococcus aureus) or secondary antibodies that will
idiomorphs The non-homologous regions of DNA, lying at the bind the Fc portions of antibodies and thereby bring them out of
same positions on homologous chromosomes, that constitute the solution.
mating type genes A and a of Neurospora. immunoaffinity chromatography Method of large-scale purifi-
idiopathic ventricular fibrillation (IVF) Ventricular fibrillation cation of an antigen (or antibody) by passing cell extracts over
without structural heart disease. columns containing immobilized specific antibodies (or antigen)
idiosyncratic drug-induced neutropenia Blood neutrophil covalently linked to an inert matrix.
deficiency occurring as an unpredictable response to a drug. immunocompetent Able to mount a normal adaptive immune
idiotope Single site (epitope) on an immunoglobulin or T-cell response against an immunogen.
receptor that is bound by a specific anti-idiotypic antibody or T- immunocompetent host An individual with a fully functional
cell receptor. One immunoglobulin or T-cell receptor usually immune system.
contains various idiotopes. immunocyte Arthropod granulocyte and/or plasmatocyte that
idiotype Collection of the idiotopes of an individual immunoglo- mediates immunological defence reactions.
bulin or T-cell receptor. immunocytochemistry Techniques by which cellular molecules
idiotypic determinant Antigenic determinant characteristic of a can be visualized by binding of labelled specific antibodies to cells
particular variable domain of an immunoglobulin or T-cell receptor. or tissues and the use of colour-based fluorescent or enzymatic
Ig See immunoglobulin. systems to detect this binding.
IHF A bacterial histone-like protein. immunoelectron microscopy Localization of intracellular struc-
IL-1 See interleukin-1. tures or of individual proteins within a macromolecular assembly
IL-2 See interleukin-2. by the binding of specific antibody bearing a label, such as gold
IL-4 See interleukin-4. particles, that can be detected using an electron microscope.

191
Glossary

immunoelectrophoresis Immunological characterization of immunoreceptor tyrosine-based activation motif (ITAM)


electrophoretically separated proteins by reaction with specific Amino-acid sequence found in the cytoplasmic portions of
antibodies. various receptors on immune system cells. It comprises ...Y-x-x-
immunofluorescence A technique in which a fluorescently [I/V]yY-x-x-[I/V]... and is a site of tyrosine phosphorylation and
tagged antibody is used to mark the cellular location of its association with protein tyrosine kinases that carry the activating
corresponding antigen (usually a protein). Fluorescently tagged signal onwards.
antigen can also be used to detect the presence of its specific immunoreceptor tyrosine-based inhibitory motif (ITIM)
antibody. Amino-acid sequence found in the cytoplasmic portions of
immunogen A substance that is able to elicit an immune response. various receptors on immune system cells. It comprises [V/L]-x-
immunogenicity The degree to which an antigen is capable of Y-x-x-L, and binds protein phosphatases that inhibit the onward
eliciting an immune response. transmission of the signal.
immunoglobulin (Ig) The class of proteins to which antibodies immunostaining See immunocytochemistry.
and their membrane-bound equivalents, the B-cell antigen immunosurveillance The constant monitoring of the body by the
receptors, belong. Single immunoglobulin molecules consist of immune system in order to prevent primary tumour development.
two identical k or l light chains and two identical a, d, e, g or m immunotherapy The attempt to augment or enhance the action of
heavy chains. They are highly variable antigen-specific the immune system response against a disease. The term often
proteins made exclusively by B cells and occur as five classes – refers to the attempt to activate the immune system to attack an
IgA, IgD, IgE, IgG, IgM – which are distinguished by their existing tumour.
heavy-chain type. Invertebrates do not possess immunoglobu- immunotrophism A theory that postulates that cells of the
lins. reproductive tract use T-cell-secreted cytokines as some of their
immunoglobulin E (IgE) One of the five classes of immuno- growth factors.
globulin. This class is usually responsible for fighting parasitic imperforate Abnormally closed.
infections. In sensitized individuals, however, IgE is the importin Also called karyopherin. The generic term for a protein
immunoglobulin responsible for much allergic disease. that recognizes a nuclear transport cargo and targets it to the
immunoglobulin fold Characteristic tertiary structure found in nuclear pore complex.
the immunoglobulin domains of members of the immunoglobu- imprinting (1) In relation to genetics, the phenomenon that for
lin superfamily. It comprises two surfaces of antiparallel b- some genes the paternal copy or the maternal copy only is
pleated sheet linked through a disulfide bridge between conserved expressed in the early embryo. (2) A developmental phenomena in
cysteine residues. which simple exposure to an object at a particular stage of
immunoisolation The separation of transplanted tissue from the development leads to a specific complex of behaviours being
immune system of the host by an artificial barrier. directed to that object at later stages of the life cycle.
immunolabelling See immunocytochemistry. in situ carcinoma Noninvasive neoplastic epithelial proliferation
immunological hypersensitivity An immune response to nor- in the breast, confined within the boundary of the basement
mally innocuous antigens by previously sensitized individuals. membrane of the terminal duct lobular unit or duct system.
Examples are allergic responses. in vitro Refers to a biological process made to occur in a laboratory
immunological memory A feature of the adaptive immune vessel or other controlled experimental environment rather than
response, in which the immune system responds more rapidly, within a living organism or natural setting.
and often more strongly, to an antigen that it has previously in vivo Refers to a biological process occurring or made to occur
encountered. within a living organism or natural setting.
immunological tolerance (1) The situation in which the inactivation Prevention of a cell, protein or gene from carrying out
immune system of an individual does not make an immune its function. A gene, for example, is said to be inactivated when it
response to a particular antigen. This may result from natural is altered so that it cannot be transcribed; an ion channel is said to
exposure to that antigen during fetal development or experi- be inactivated when it no longer conducts ions although it
mental exposure to high doses later in life. (2) The acceptance of a remains open.
transplanted organ without continuing immunosuppressive drug inbreeding Self-fertilization in hermaphrodite individuals or, in
therapy. outcrossing species, mating among individuals that are geneti-
immunology The study of the immune system. cally closely related, such as brother–sister matings.
immunomodulation An increase or decrease in the level of inclusion body (1) Granular proteinaceous body in cytoplasm. In
immune response brought about by various specific and bacteria genetically engineered to express a recombinant protein,
nonspecific means. they often contain a high content of the protein. (2) A localized
immunopathology Damaging effects caused by an immune accumulation of viral protein in infected cells. May have
response, such as tissue damage due to overly aggressive cytotoxic distinctive appearance, location or stain uptake properties.
lymphocytes and autoreactive antibodies (autoimmunity) incoming solar radiation See insolation.
immunophenotype The pattern of gene expression manifested by incompatibility The impossibility of two different plasmids
any cell population with specific reference to proteins, or coexisting in the same cell. Incompatibility results from cross-
carbohydrates bound to proteins, which can be detected by reaction between replication control systems and, to a lesser
antibodies. extent, partition mechanisms.
immunopotentiator A substance capable of raising an immune incompatibility group A group of plasmids whose replication
response. control systems are sufficiently closely related that they inhibit
immunoprecipitation Precipitation of an antigen from a soluble each other’s replication. Used as a convenient method of plasmid
cell extract by binding to a specific antibody. classification.

192
Glossary

incubation period The time from contracting an infectious agent inhibitory postsynaptic potential (IPSP) An alteration in
to the first clinical symptoms of infection. membrane potential in a post-synaptic cell (neuron or muscle),
incus Small bone in the mammalian middle ear that articulates with as a result of a stimulus such as a neurotransmitter, which inhibits
the stapes and the malleus. It is homologous with the quadrate of the cell from producing an action potential.
nonmammalian amniotes. In many Mesozoic forms it is a inhibitory synapse A synapse that when activated inhibits the
substantial bone directly articulated to the jaw. postsynaptic neuron from producing an action potential.
indehiscent Describes a fruit that does not open by splits or pores. initial lymphatic Microscopic lymph channels made up of
indigenous Living naturally in the region. endothelial walls without a smooth muscle coat. Previously
individual In ontology, a general term for any concrete, particular called terminal lymphatics or lymphatic capillaries.
thing whatsoever, not just an organism. initial segment That part of the axon closest to the cell body. It is
induced systemic resistance Disease resistance in plants due to a unmyelinated and extends to the start of myelination.
physiological change in the plant that is induced by nonpatho- initial velocity Rate of unidirectional transport across a membrane
genic organisms. measured before any back-flux of substrate occurs.
inducible enzyme An enzyme that can be induced in response to a initials Cells in plant meristems that remain undifferentiated and
stimulus. continue to divide.
induction of differentiation The dependence of differentiation in initiation The first step of translation and protein synthesis, where
one tissue on signals from other tissues. It serves to correlate the initiator tRNA (Met-tRNAI) is correctly matched to the
differentiation with other developmental and functional events. AUG start codon in the mRNA in the ribosomal P site.
induction (1) A cellular interaction in embryonic development in initiation codon A codon specifying the first amino acid of a
which signals from one cell (or group of cells) cause another cell polypeptide chain (e.g. AUG).
(or group of cells) to change its developmental fate. (2) The initiation factor IF-2 Prokaryotic initiation factor that ensures
production of a specific protein following activation of its gene in correct ribosomal P-site binding of the initiator fMet-tRNAfMet.
response to a particular signal, often environmental. initiation factor IF-3 Prokaryotic initiation factor that carries out
inductive signals In development, signals delivered by extracel- proofreading of the 30S ribosomal initiation complex.
lular molecules to bring about cell differentiation and migration. in-line attack Attack of an activated nucleophile (e.g. OH) on
Cells release inductive signals to organize development. phosphorus ‘in line’ with the phosphodiester bond to be cleaved.
infection Invasion and replication within a host organism by innate immunity Immediate antimicrobial responses due to pre-
microorganisms. existing components of the immune system whose intrinsic
infectious clone Complete complementary DNA (cDNA) copy activity is unaffected by, and not specific for, a particular antigen.
of viral RNA that can be used to obtain infectious transcripts innate responses See innate immunity.
in vitro. The cDNA copy can also be incorporated in a inner cell mass (ICM) Portion of the mammalian blastocyst that
plasmid, which upon entering the bacterial cell leads to phage will form primitive endoderm and the epiblast.
infection. innervate See innervation.
infective centre An entity in a phage-infected cell culture that is innervation Establishing contact of a nerve with its target tissue.
capable of forming a single plaque. It is either a phage particle or innovation See novelty.
an infected cell that has not yet lysed when plated. inoculum A small number of individual cells used to initiate a
inferior Towards the feet. bacterial culture.
infertility In humans, the inability to conceive after one year of iNOS See inducible nitric oxide synthase.
unprotected intercourse. inosine A purine base similar to adenine, but lacking one amino
inflammation Infiltration of leucocytes and plasma fluid and group. Present in RNA in small amounts, it base pairs with
proteins into extravascular tissues, causing swelling, pain, cytosine instead of uracil.
reddening and heat. It can be caused by infection or tissue inositol 1,4,5-trisphosphate (IP3) An intracellular signalling
damage and, in the case of infection, both initiates and is molecule that is generated by the hydrolysis of membrane
amplified by a subsequent immune response. phospholipids via activation of a G-protein-coupled phospho-
inflorescence The shoot of a plant during the reproductive phase lipase. IP3 triggers the release of calcium from intracellular stores.
of the life cycle, bearing the flowers or other reproductive INR See International Normalized Ratio.
structures. insertion element A mobile DNA sequence in bacteria that
infradian rhythm A rhythmic process within an organism with a carries only the genes needed to move it from one site to another in
phase longer than 24 hours. DNA. Its insertion into another site generally causes a mutation.
infundibulum A funnel-shaped structure lying in ctenophores insertion mutagenesis See insertional mutagenesis.
immediately aboral to the stomodaeum, corresponding to the insertional inactivation See insertional mutagenesis.
stomach or oesophagus of higher invertebrates. insertional mutagenesis Disruption of gene function by the
infusion A culture made by soaking leaves or grasses in water, with insertion of a foreign DNA sequence into a gene.
or without moderate heating, and then allowing the microbial inside-out signalling Signalling from within a cell that leads to
community to grow. changes in integrin-mediated cell adhesion.
infusion-related adverse effect An acute response to intravenous insolation The radiant energy received on Earth from the Sun. It is
infusion of an agent. currently 1350 watts m2 at the top of the atmosphere, at the
inguinal Pertaining to the groin area. Equator. Acronym for incoming solar radiation.
inhibitory neurotransmitter Chemical released from nerve insonation The exposure of a region of tissue to ultrasound.
terminals that produces inhibition of a target neuron. instar The period between two moults in arthropods.

193
Glossary

integral membrane protein Membrane protein that interacts intermediate host A host in which no sexual reproduction of a
strongly with the lipid bilayer and can be solubilized only by parasite occurs.
detergents. Such proteins have a portion of their mass buried in intermediates Short-lived chemical species hypothesized as
the lipid bilayer. occurring transiently on the way from reactants to products
integrase The transposase encoded by a retrovirus that mediates in chemical reactions.
the integration of the DNA form of the virus into chromosomal internal state Enzyme–ligand or enzyme–transition state com-
DNA by site-specific recombination. plex.
integration host factor (IHF) Bacterial protein required for the International Normalized Ratio (INR) Correction of the pro-
integration of phage l into the host cell genome. thrombin ratio – the patient’s prothrombin time : control
integrins A superfamily of heterodimeric cell adhesion molecules prothrombin time (PT) – by the International Sensitivity Index
that mediate cell–cell and cell–matrix interactions, and thereby (ISI) assigned to a thromboplastin, calculated as follows: INR 5
control multiple cellular responses. They each consist of one a (PT ratio) ISI.
and one b subunit. Internet Worldwide digital communications network, conceived
integron A genetic cassette within which antibiotic-resistance genes and developed originally by the US Department of Defense and
are acquired, lost and reshuffled by site-specific recombination. now transferred to international commercial enterprises.
integumentary structures Structures related to the epidermis (skin). interneuron Neuron that is interposed between, and synaptically
integument Skin. connected with, two other neurons.
intercalary regeneration A type of regeneration that fills space interphase The time a eukaryotic cell spends between successive
between the edges of a wound. mitoses. There is considerable variation in duration of interphase
interclavicle A midline T-shaped bone present in the ancestors of between individual cells within an actively dividing population.
the mammals and in early mammals. It connects the right and left interspecies hydrogen transfer Production and consumption of
pectoral girdles. Among extant mammals it is present only in hydrogen (H2) by different groups of microbes interacting closely
monotremes. during anaerobic catabolism.
interdigitating dendritic cell Large antigen-presenting cell with interspecific Between two or more species.
extensive and numerous processes found in T-cell areas of interspersion pattern The pattern in which repeated sequences
peripheral lymphoid organs. and single-copy sequences are intermingled in the genome.
interfascicular Refers to the area between vascular bundles. interstitial (1) Pertaining to or belonging to interstices or
interferons A family of proteins with antiviral activity. interspaces of a tissue or organ. (2) Pertaining to the spaces
interfertile Able to exchange genes. between mineral grains.
intergenic recombination Recombination between closely linked interstitial cell Stem-cell population in cnidarians giving rise to
homologous genes, following chromosomal misalignment at nerve cells, gland cells, nematoblasts and nematocytes (stinging
meiosis. The recombination event may be crossing-over or gene cells), and gametes (observed only when the animals follow the
conversion, and results in a gene that represents a hybrid of the sexual cycle).
two misaligned genes. interstitial organism Organism that inhabits the spaces between
interleukin-1 (IL-1) A cytokine produced by macrophages and sand grains.
epithelial cells that synergizes with other factors to stimulate intersubunit space Partially open space between the small and the
colony growth from bone marrow. large ribosomal subunits, whose shape is tailored to accommo-
interleukin-2 (IL-2) Cytokine secreted by activated T cells that date tRNAs.
stimulates the growth of T cells. intertidal Describes the area covered by the sea at high tide and
interleukin-3 (IL-3) Cytokine produced by T cells and epithelial exposed at low tide.
cells, also known as multi-colony-stimulating factor, this intervening sequence See intron.
stimulates the growth of multiple types of colonies from bone interventional radiology The use of an imaging procedure to
marrow stem cells, i.e. blast cell, erythroid, megakaryocyte and control an interventional procedure, as distinct from making a
mixed colonies. diagnosis.
interleukin-4 (IL-4) Cytokine secreted by T cells and mast cells intima An inner layer of cells in the pulmonary vascular wall,
that stimulates B-cell activation and isotype switching. consisting of cells of endothelial or perhaps smooth muscle
interleukin-5 (IL-5) Cytokine produced by T cells and mast cells derivation.
that stimulates the growth of eosinophils. intracellular messenger Any intracellular ion or protein pro-
interleukin-6 (IL-6) Cytokine required for bone marrow stem cell duced as a result of receptor stimulation and which directly
growth in vivo. It stimulates the production of granulocyte and modifies the activity of another protein.
granulocyte–macrophage colonies from bone marrow cells in intracytoplasmic membrane The photosynthetic apparatus in
vitro and the final stages of B-cell differentiation. It also has purple phototrophic bacteria.
effects on the liver and bone formation. intramembranous ossification Bone formation and repair
interleukins Members of the cytokine family that were originally achieved through the direct transformation of mesenchymal
thought of as intercellular messengers between leukocytes but are stem cells into osteoblasts, without requiring the intermediate
now perceived as having wider immunological and inflammatory step of a cartilage anlage.
effects. in trans On or affecting another molecule.
intermediate filaments Protein filaments of the eukaryotic cell intraspecific Within a species.
cytoskeleton intermediate in diameter between microtubules and intraspecific variation Variation, either morphological or genetic,
microfilaments. They are constructed from a family of related that occurs within a species so that no two individuals are exactly
intermediate filament proteins. alike unless they are clones.

194
Glossary

intrastrand exchange Recombination between copies of a island model migration The random movement of individuals
tandem array of sequences on the same DNA strand. between populations.
intrinsic rate of population growth The instantaneous rate of isoacceptor tRNA A member of a set of tRNAs that accept the
growth of a population not subject to density limitation. Usually same amino acid.
symbolized as r. isochromosome A structurally abnormal chromosome composed
introgression The transfer of genes between genetically differ- of either identical short or long arms, formed by the aberrant
entiated populations or species. division of the centromere.
intromittent organ The male copulatory organ. isodiametric Of similar dimensions in all directions.
intron A noncoding region in an RNA (or a gene) that is removed isoenzymes Enzymes with identical substrate specificity but of
from the initial RNA transcript by splicing during maturation to different biochemical structure.
produce the functional RNA. Also known as an intervening isoforms Multiple forms of a molecule (e.g. a protein) that perform
sequence. the same function (e.g. catalyse the same reaction in the case of
intron phase The position of an intron within the reading frame of isoenzymes) but differ in their composition (i.e. amino-acid
the coding sequence. sequence in the case of a protein).
intronless Describes a gene that is made up of a single long exon isogamontic ciliates Ciliate protozoans in which the co-conju-
and thus lacks introns. gants do not differ from each other morphologically.
invariant lineage Describes the developmental condition in which isograft A tissue or organ transplant from one individual of an
the corresponding cells in different individuals always have the inbred strain to another. Such grafts not rejected.
same descent. Cell fates are thus invariantly specified. isomerase An enzyme that catalyses the interconversion of two
invasion The spread of tumour cells to surrounding tissues. isomers, i.e. two compounds comprising the same atoms.
invasive species A species introduced from its native range to a isomerization The process of interconverting isomers, i.e. differ-
new area (also known as exotic or introduced species). Invasive ent covalent arrangements of the same atoms.
species can compete with or prey on local species, at times causing isometry The situation when there is a linear proportionality in
extinctions and shifts in community composition and function. respect to increase in size, e.g. a tenfold increase in mass and
inversion Structural rearrangement within a chromosome result- volume of heart, muscle, blood, and lung is associated with a
ing from two breaks in the chromosome with inversion of the tenfold increase in body mass.
intervening segment of chromosome before rejoining. isomorphous replacement A technique in X-ray crystallography
invertebrate Any of the animals without vertebrae or backbones. in which a heavy metal ion is introduced into a crystal without
All animals other than vertebrates. changing the crystal’s shape, in order to determine the signs
ion channel A transmembrane protein that forms a selective pore (positive or negative amplitudes) of diffracted X-rays.
for the diffusion of ions through a cell membrane. Ion channels isoschizomers Restriction enzymes from different bacterial
are usually specific for a certain type of ion and most are gated, in species that recognize and cleave identical DNA sequences.
which case they open only in response to a particular stimulus isotope signature The evidence that organic molecules have been
such as a change in membrane potential or the binding of an processed by living systems, which can be obtained by analysis of
extracellular ligand. isotopes of e.g. carbon.
ionic strength A measure of the total ionic concentration of a isotopes Physically and chemically different atoms of the same
solution regardless of net charge. element, resulting from differences in the number of neutrons in
ionotropic receptor A ligand-activated ion channel. When ligand the atomic nuclei.
molecules (typically neurotransmitters) bind the extracellular isotopic tracer technique A method of investigating biosynthetic
portion of the receptor, the receptor channel opens and allows mechanisms by feeding an organism with a putative biosynthetic
ions to flow through the channel and across the cell membrane. intermediate labelled with an isotope not naturally present in
IP3 See inositol 1,4,5-trisphosphate. large amounts (e.g. 13C, 14C, 2H, 3H, 15N, 18O).
ipsilateral Belonging to or occurring on the same side of the body, isotopomer A molecule that has been labelled, in a known
as opposed to contralateral, which refers to the opposite side of position, with an isotope (distinguishable by nuclear magnetic
the body. resonance) of the element normally found at that position.
IPSP See inhibitory postsynaptic potential. isotype The class of an antibody (e.g. IgG, IgA, IgM), which
iridium An element in the platinum group having the atomic depends on the type of its heavy chain.
number of 77. Iridium is enriched in some extraterrestrial isotype switch The change in a B cell or its progeny from the
asteroids, and is depleted in the Earth’s surficial rocks. secretion of immunoglobulin (antibody) of one class to immu-
iridocyclitis Inflammation of the iris and ciliary body. noglobulin with the same V regions but a different heavy-chain C
iron reduction A form of anaerobic bacterial decay in which iron region and, hence, a different isotype (or class).
hydroxides or oxides serve in the metabolism of organic matter. isoxazolylpenicillins A group of semisynthetic penicillins that are
iron–molybdenum cofactor (FeMo-co) The active site cofactor not inactivated by staphylococcal b-lactamase. They include
of dinitrogenase: Fe7S9Mo:homocitrate. oxacillin, cloxacillin, dicloxacillin and flucloxacillin.
ischaemic heart disease A disease of the coronary artery wall isozymic forms Physically distinct forms of an enzyme present in
characterized by atheromatous plaque formation reducing blood different cells.
flow to the myocardium. ITAM See immunoreceptor tyrosine-based activation motif.
ischial callosities Hard pads present on the ischial bones of iteroparity Repeated episodes of reproduction within the life cycle.
monkeys and partially on gibbons, an adaptation for sitting on ITIM See immunoreceptor tyrosine-based inhibitory motif.
branches. IVF See idiopathic ventricular fibrillation.

195
Glossary

Janus kinase (JAK) A member of a family of tyrosine kinases that keratinocyte The predominant cell within the epidermis, contain-
includes the interferon-associated enzymes Tyk2 (tyrosine kinase ing large amounts of the protein keratin.
2), JAK1 (Janus kinase 1) and JAK2 (Janus kinase 2). They act by keratins The protein subunits of certain cytoskeletal intermediate
activating STATs which then act as transcription factors. filaments. They interact with integrins, thereby contributing to
Jarisch–Herxheimer reaction A transient, short-term immuno- cell strength and to cell motility during migration.
logical reaction, commonly seen after antibiotic treatment of ketogenic Describes metabolites that result in the production and
syphilis and sometimes other diseases, due to liberation of excretion of ketone bodies.
endotoxin or antigens from the killed microorganisms. It is ketone body Acetoacetate, D-b-hydroxybutyrate or acetone,
marked by fever, chills, headache, muscular and cutaneous which are water-soluble derivatives of fatty acids produced by
lesions. the liver during starvation or in uncontrolled type I diabetes.
jasmonic acid Plant hormone, a structural analogue of animal key innovation A novel character that may have been important in
prostaglandins, required for wound-induced gene expression. the origin of an adaptive radiation.
jaundice A yellow appearance resulting from increased bile Kimmeridgian The middle stage of the Upper Jurassic.
pigment in the plasma. kin selection Selection that results from close kin behaving in ways
joint molecule An intermediate in the DNA recombination that favour each other, resulting in increased inclusive fitness.
process consisting of single-stranded DNA paired to a homo- kinase An enzyme that catalyses the transfer of the terminal g-
logous region within a double-stranded DNA molecule. phosphoryl group of ATP to an acceptor molecule, with the
Jurassic period The middle major division of the Mesozoic era. formation of ADP and the phosphorylated acceptor. Protein
just-so-story An adaptationist explanation for a trait that relies on kinases are enzymes that transfer phosphoryl groups in this way
plausible but generally untestable claims about the circumstances to proteins, thus phosphorylating them at specific sites and often
under which the trait is supposed to have evolved. A just-so-story changing their activity as a result.
is a ‘how possibly’ explanation as opposed to a ‘how actually’ it kinematics The study of the characteristics of motion.
happened. kinesin A member of a superfamily of motor proteins with ATPase
activity that are present in most eukaryotic cells. They associate
with microtubules and move along them, hydrolysing ATP to
produce the energy to power the movement.
K antigen Bacterial capsular antigen used for serotyping.
kinesis Mobility of a structure (of a skull, typically) about
K homology (KH) motif An RNA-binding domain in proteins
particular joints.
that consists of ~60 amino acids and contains a conserved
kinetics (1) A branch of chemistry that deals with the study of rates
octapeptide, Ile-Gly-X2-Gly-X2-Ile, where X varies among the
and rate laws of chemical reactions. (2) The study of how forces
different KH motifs but is usually a positively charged amino
act on a system to create movement.
acid.
kinetochore Specialized structure that forms at the centromere
kairomone (1) A substance produced by a predator that induces an
and to which the microtubules attach during mitosis and meiosis.
adaptive change in the development of the prey. (2) Plant
kinetoplast Organelle containing mitochondrial DNA that char-
chemical that attracts herbivores.
acterizes the protozoan order Kinetoplastida.
kala-azar Visceral leishmaniasis.
kinetosome Subpellicular, cylindrical structures composed of nine
kaliuresis Increased potassium in the urine. longitudinal equally spaced peripheral tubules each composed of
Kaposi’s sarcoma-associated herpesvirus A herpesevirus asso- three microtubules. The embedded basal body produces a cilium
ciated with the vascular cancer, Kaposi’s sarcoma. or flagellum.
karyogamy Fusion of two nuclei (usually haploid nuclei). kinety A row formed by kinetosomes, the basal bodies of cilia.
karyogram Representation of an entire chromosome set that has Kingdom In taxonomy, the second highest level of biological
been stained by one of several possible methods to yield discrete classification.
banding patterns. kissing complex Complex formed by two stem–loops in a nucleic
karyonide In ciliate protozoans, a vegetative cell lineage that is acid base pairing in their respective loop regions.
derived from an exconjugant cell with a single newly formed Klenow fragment The fragment of Escherichia coli DNA poly-
macronucleus. merase I derived by removal of the 50 –30 exonuclease domain. It
karyopherin See importin. has the function of adding bases to the unpaired stretch of DNA
karyotype A complete description of the number and morphology at the end of a growing DNA duplex.
(often a photograph) of all the chromosomes in a cell as they kleptoplastidy The capacity of certain microorganisms to use the
appear at mitosis or meiosis. chloroplasts of the prey they ingest in order to photosynthesize.
kb Kilobase, a unit of measurement of the length of nucleic acid Km The Michaelis constant. A kinetic constant related to Ks, and
molecules (DNA or RNA). It is 1000 nucleotides or base pairs. which describes the concentration of substrate required to half-
kcat The first-order rate constant that describes the rate of an saturate the enzyme molecules during steady-state catalysis.
enzyme-catalysed chemical transformation of substrates to KNM-ER Abbreviation of the prefix ‘Kenya National Museum –
products. East Rudolf’.
kcat/Km Parameter that describes the catalytic efficiency of an knockout mouse A mouse in which a given predetermined gene
enzymatic reaction involving a particular substrate. The larger has been inactivated. It is generated by first targeting the gene in
the value, the more efficient the reaction for a particular enzyme– cultured embryonic stem cells and then introducing the altered
substrate pair. stem cells into a blastocyst.
keratinized epithelium The hard, dry, ‘horny’ outer surface knot A closed circular DNA including a knot which cannot be
found on skin and related structures. removed without double-strand breakage.

196
Glossary

Kranz anatomy The wreath-like arrangement of bundle-sheath laser ablation Experimental technique used on C. elegans in which
cells and mesophyll cells within the leaves of C4 plant species. a single cell is killed by means of a microscopically focused laser
Krebs cycle The tricarboxylic acid cycle, a metabolic pathway beam.
within mitochondria by which carbohydrates and fats are fully late genes Viral genes transcribed late in the replication cycle,
combusted during aerobic respiration. mainly virion proteins.
krill Large schools of planktonic euphausiacean crustaceans (sub- latency The activity of a mitochondrial enzyme following the
class Eumalacostraca) typically found in Antarctic waters. Also disruption of mitochondria minus its activity in the intact
refers to the euphausiaceans themselves. preparation. Enzyme latencies are used to assess the integrity of
Ks The equilibrium dissociation constant for the binary complex mitochondria.
between an enzyme and its substrate molecules. latent period (1) The time needed for a vector that has completed
K-shell The innermost shell or orbital of electrons in an atom. the acquisition of virus to become infective. (2) The entire period
K-shell binding energy The energy required to expel an electron during which phage reproduction is occurring. It comprises the
from the K-shell of an atom. interval between adsorption of phage to the cell and lysis of that
cell.
Kyte and Doolittle hydropathy analysis Method of assessing the
lateral Away from the midline.
hydrophobicity of amino acids in a protein. This information is
lateral diffusion Movement by lipids and by proteins in the plane
used to predict membrane-spanning domains and topology.
of a membrane.
lateral geniculate nucleus A nucleus in the thalamus of the brain
that relays visual information from the retina to the primary
labelling index In studies of actively growing and dividing cells, visual cortex. Additional inputs to the lateral geniculate nucleus
the percentage of cells labelled with tritiated thymidine from the brainstem and primary visual cortex modulate its
([3H]TdR), which reflects DNA replication, is known as the output.
labelling index. lateral inhibition The process whereby a developing cell inhibits
labyrinth The inner layers of the placenta in rodents. its neighbours from assuming a similar fate.
lacteal Lymph vessels of small intestine. lateral root Plant root formed on a young root that has not yet
LAG See line of arrested growth. started secondary growth.
Lagerstätte (plural Lagerstätten) A fossil deposit of exceptional Laurasia The northern continental landmass formed during the
preservation. This is a specialized use of a German word that early Mesozoic. It included what were to become North America,
originally referred to a rich mineral deposit. Europe and Asia.
lagging strand In a replicating DNA, the strand that is Lazarus taxa Taxa that disappear from the geological record, only
synthesized discontinuously and in the opposite direction to fork to reappear at a later time.
movement, as a series of short Okazaki fragments. leading strand In replicating DNA, the strand that is synthesized
lambdoid bacteriophages The family of related phages of which continuously and in the same direction as the fork moves during
l is the prototype. DNA replication.
lamella See bilayer Leäo spreading depression Depression of nerve cell function in
lamellar bone Bone in which fine collagen fibre bundles are selected areas of the brain causing the aura of migraine. It also
arranged in thin plates, or lamellae, with different orientations in affects cranial blood vessels.
successive lamellae. leaving group Structural entity produced by cleavage of a
chemical bond, requiring stabilization (e.g. by protonation)
lamellated bone Bone that has a gross layered structure due to
before release of the product.
variation in the mineral structure.
lecithotrophic Describes the nourishment of larva solely by
lamellipodia (singular lamellipodium) Actin-based motile veil-
digesting yolk.
like protrusions from the leading edge of a migrating eukaryotic
lectin A class of proteins first found in plants, but also present in
cell, which assist in locomotion.
animals, which bind to sugars on e.g. the glycoproteins and
laminar Sheet-like. glycolipids of the plasma membrane. They are often used as
lamivudine Nucleoside analogue inhibitor of human immuno- receptors in innate defence reactions. Lectins found in arthropods
deficiency virus (HIV) reverse transcriptase. Used as an anti-HIV are known as haemagglutinins or heteroagglutinins.
drug. LEF An HMG-box transcription factor that binds DNA with
Lancefield groups A serotyping scheme (devised by Rebecca sequence specificity, altering the bending of DNA and thus gene
Lancefield) for streptococci, based on antigenic cell wall expression.
polysaccharides. Grouping is usually accomplished by precipita- legume Fruit (a dehiscent pod) of the Leguminosae (peas, beans,
tion reactions with group-specific antisera. vetches, etc.). The term is also used to describe the plants of this
landrace A local or regional crop variety that has arisen from farm- family.
level seed production and selection, often over centuries. Leishmania Genus in the Trypanosomatidae family of protozoa.
Langerhans cells Immature dendritic cells of epithelium, particu- lek Mating arena where males display and females come simply to
larly the skin, which pick up antigen and carry it to draining acquire matings. No resources are associated with a lek besides
lymphoid organs. males.
lariat The looped structure formed in the intron as a result of the lens The transparent structure in the eye that focuses the optical
formation of a 20 –50 phosphodiester bond between the first base image to the retinal photoreceptor layer. Its shape ranges in
of the intron and an adenosine (branchpoint adenosine) in the different animals from almost spherical to a flat disc with an
first step of splicing. ellipsoid section.

197
Glossary

lentic Pertaining to slow moving water characteristic of a pond or Limulus lysate test An assay to detect the presence of bacterial
lake. lipopolysaccharide based on the ability of this molecule to cause
leptospiraemia Presence of leptospire bacteria in the blood. gelation of amoebocyte lysate of the horseshoe crab, Limulus
leptospiruria Presence of leptospire bacteria in the urine. polyphemis.
Ler The Arabidopsis ecotype Landsberg erecta. line of arrested growth (LAG) A line that marks a pause in bone
lesion A site of damage. In DNA, the term refers to damage such as growth.
an adducted base or a strand break. lineage (1) A sequence of fossil species, which usually do not
leucocytes A general term for all white blood cells. Leucocytes overlap in time, but which form part of a distinctive evolutionary
include lymphocytes, monocytes, dendritic cells and polymorpho- trend. (2) Descendants from a particular precursor cell.
nuclear lymphocytes. linear energy transfer Energy deposited in material per unit of
leucocytosis Increased count of white blood cells in the blood track by a transmitting ionizing particle (usually expressed as
circulation. keV mm1). It can be regarded as a measure of the efficiency of a
leukocyte See leucocyte. radiation for producing a biological effect.
Lewis acid An electron pair acceptor. Metals often serve as Lewis linkage The situation when two loci are situated close together on
acids in catalysis by accepting an electron pair from a substrate (to the same chromosome, so that their alleles usually stay together at
form a coordinate bond), thereby polarizing the electron density meiosis in gamete formation.
of the substrate. linkage disequilibrium Nonrandom association between alleles at
different genetic loci such that they tend to be inherited together.
library Collection of cloned DNA fragments. Libraries can be
It is due either to physical proximity on the chromosome or to
composed of genomic DNA clones, generated from fragments of
selection for a particular combination of traits.
genomic DNA, or of cDNA clones, generated from reverse
linker DNA The DNA joining two nucleosomes, which is lost when
transcription of cellular mRNAs.
a nucleosome is trimmed to a core particle.
life cycle Refers to various developmental stages that
lipid Substance insoluble in water but soluble in organic solvents
one organism, or even one cell, passes through during
such as chloroform and acetone. Lipids include phospho- and
its life. Some parasites exhibit life cycles involving several
glycolipids, sterols, waxes and triacylglycerols.
hosts.
lipid bilayer The structure formed by amphipathic phospholipids
lifetime In regard to a fluorophore, the average amount of time
in an aqueous environment in response to the hydrophobic effect.
between light absorption and emission.
The molecules are organized in a double layer in which the polar
ligand (1) The smaller of two molecules that interact reversibly to
headgroups face outwards (to the aqueous environment) and the
form a complex. (2) A molecule that binds to a cell-surface
hydrocarbon chains are in the interior. A lipid bilayer is the basic
receptor. (3) A molecule or ion that binds to a metal ion by
structure of all biological membranes.
donation of an electron pair.
lipid monolayer A planar array of amphipathic lipid molecules,
ligase An enzyme that joins two molecules together covalently, e.g. one molecule thick, where the hydrophobic surface is formed by
D-alanine ligase, which forms D-alanyl-D-alanine from two D-
the lipid hydrocarbon chains and the hydrophilic surface is
alanine molecules. established by the polar headgroups.
light-harvesting complex A protein complex that absorbs light lipophilicity A molecular property associated with an affinity for
energy and converts it to exciton energy that can migrate to a oily or waxy environments as opposed to those that are aqueous
neighbouring complex or to a reaction centre. The light is in nature.
absorbed by pigment molecules (e.g. chlorophyll and carote- lipopolysaccharide (LPS) A component of the cell walls of Gram-
noids) that are attached to the protein. negative bacteria consisting of a lipid A moiety and a highly
light-harvesting structure The main light-gathering antenna species- and strain-specific polysaccharide domain. The verte-
(complexes of chlorophyll and protein) in photosynthetic brate innate immune response has evolved to be exquisitely
organisms. sensitive to minute amounts of LPS.
lignocellulose The mixture of polymers found in the cell walls of lipoprotein A complex of proteins and lipids that transports lipids
higher plants. It is composed of cellulose microfibrils associated in the blood.
with smaller, more complex polysaccharides called hemicellu- liposome A tiny synthetic lipid vesicle into which protein or other
loses embedded in a matrix of lignin. The adjective is materials, e.g. fluorescent molecules, can be incorporated for
lignocellulosic. delivery into cells.
likelihood A general method of testing hypotheses that contrasts liquid crystal The type of organization within the lipid bilayer
the exact probabilities of observing data given particular membranes of cells. There is extensive order in the plane of the
hypotheses. membrane. Within this nearly two-dimensional structure, lipid
likelihood-ratio statistic A statistic based on the ratio of two molecules can diffuse laterally in two dimensions but experience
probabilities, used to test the hypotheses under which the very little freedom to move between layers.
probabilities were generated. liquid scintillation Process by which radioactivity in a sample can
limb plexus Region at the base of the developing limb where be quantified through the use of a solvent (a fluor) that has the
the incoming axons join together and sort out from one ability to absorb energy and re-emit it as light.
another. lithification The alteration or hardening of loose sediment into
limestone Rock composed mostly of calcium carbonate. compact, cemented rock.
limited proteolysis Proteolytic hydrolysis of an entire organelle or lithosphere The upper layer of the Earth’s crust, which moves in
membrane-bound protein(s) and cleavage of surface-accessible many separate tectonic plates across the weaker, plastic astheno-
peptide regions. sphere below.

198
Glossary

lithostratigraphic unit A body of rock with certain unifying lymphadenopathy Pathological enlargement of one or more
features that enables it to be recognized, mapped and correlated lymph nodes.
as a single unit. lymphangiogram The detection by radiology of contrast medium
littoral The seashore, or, living on the seashore. injected into the lymphatic system, used to highlight abnormal-
lobopod A limb constructed as a lobe-like outgrowth of the body ities of that system, including lymph nodes.
wall in tardigrades and lacking rigid sections and articulations. lymphangion A collecting lymphatic segment with an upstream
local alignment The alignment of two or more sequences at and downstream valve, and intrinsic smooth muscle to compress
discrete regions of similarity. This type of alignment is well suited the lymphatic lumen.
for database searching with an uncharacterized query sequence. lymphatic system The lymphatic vessels and lymphoid tissues.
local response Activation of wound-response genes in damaged lymphatic vessels, lymphatics Vessels that carry the lymph
tissues of a plant. Usually refers to responses occurring in the leaf formed in various tissues to lymph nodes and eventually from
that has actually been wounded. there to the blood circulation.
locomotion Movement of a cell or organism. lymphocyte The cell type mediating the antigen-specific part of the
locus Position on a chromosome at which the gene controlling a adaptive immune response. It is a type of white blood cell with a
particular trait resides. It can be occupied by different alleles of large, spherical nucleus surrounded by a thin layer of non-
the gene. granular cytoplasm. The two main classes of lymphocytes are the
locus heterogeneity See genetic heterogeneity. T lymphocytes (or T cells) and the B lymphocytes (or B cells). B
loess Wind-blown silt derived from deserts or from the rock-flour cells develop into antibody-secreting cells, while T lymphocytes
produced by glaciers. become either cytotoxic T cells or helper T cells.
long QT syndrome (LQT) A cardiac disorder characterized by lymphocyte homing Migration of lymphocytes that directs them
prolongation of the QT interval on electrocardiograms, syncope to particular tissues where they may develop further and/or carry
(sudden loss of consciousness), seizures, and sudden death due to out their effector function.
a specific ventricular tachycardia called torsade de pointes. lymphocyte recirculation The process in which naı̈ve lympho-
long-branch attraction The tendency of parsimony-based cytes are continuously migrating from the bloodstream into
methods to group long-branch taxa together. secondary lymphoid organs and returning to the blood via
longevity The ability of seeds to maintain their viability over a long efferent lymphatics.
period. lymphoedema Oedema resulting from lymphatic obstruction.
long-term potentiation (LTP) A long-lasting (41 hour) increase lymphoid cells Lymphocytes or their precursors.
in synaptic strength caused by a brief period of synaptic activity lymphoid follicle An accumulation of rapidly proliferating B
that fulfils special requirements. For many forms of LTP, this lymphocytes in a secondary lymphoid tissue.
requirement is that the postsynaptic cell be strongly depolarized. lymphoid progenitor A cell that is committed to a developmental
look-up table (LUT) An array of values that changes a grey value pathway that will result in lymphocytes.
of a pixel to another value. lymphokines Cytokines produced by lymphocytes.
lophophore Tentaculated extension of the mesosome that lymphoma A cancer of lymphoid tissue.
embraces the mouth, but not the anus, in phoronids. Its main lymphopenia A deficiency of lymphocytes in the blood circula-
functions are feeding, respiration and protection. tion.
lorica A loosely fitting secreted extracellular shell that can partially lymphoproliferative A disease or disorder that results in a
or nearly completely envelop the cell body of suctorian protozoa. population of lymphocytes that proliferate in an uncontrolled
lotic Pertaining to rapidly moving water characteristic of a river or manner.
stream. lymphotoxin Cytokin produced by activated T cells and which is
low-threshold channel A voltage-gated channel that opens when toxic to some cell types. Also known as tumour necrosis factor-b
the membrane potential is depolarized only a few millivolts above (TNF-b).
resting potential. lyophile A sample that has been preserved by freeze-drying.
LPS See lipopolysacharide. lyriform organ A sensory organ unique to arachnids that is formed
LQT See long QT syndrome. by several closely spaced slit sensilla and is used in detecting
LTD See long-term depression. compressional forces in the exoskeleton.
LTP See long-term potentiation. lyse To rupture a cell, releasing its contents and resulting in cell
lumen (1) The interior space within a hollow structure such as the death.
gut or a blood vessel. (2) The water phase within the thylakoid lysis Rupture of a cell, resulting in the release of its contents and cell
vesicle of chloroplasts. death.
luminal Pertaining to a lumen, or facing into the lumen. lysis inhibition A delay in the metabolic death of a phage-infected
lupus erythematosus An autoimmune disease of unknown cell, with consequent delay in lysis, caused by infection of a
aetiology with disseminated proliferative lesions of minute blood previously infected cell.
vessels and lymphatics. lysogen A bacterium that harbours a phage chromosome
LUT See look-up table. in a stable inactive association with the bacterial chromo-
Lyapunov exponent A measure of whether nearby trajectories some.
will tend to converge (negative), diverge (positive) or retain their lysogenic Describes a type of bacteriophage that can integrate into
separation (zero). A positive Lyapunov exponent is a hallmark of the host DNA and remain inactive there for many cell
chaos. generations. See lysogeny.
Lyme borreliosis See Lyme disease. lysogenization The establishment of a lysogen.

199
Glossary

lysogeny A stable state in the life cycle of a temperate phage, in macroprojectile Any macroscopic object that can be accelerated
which the phage genome is integrated into the host cell to high velocity, which can thereby accelerate the microcarriers
chromosome in an inactive state (prophage), with no synthesis placed on its surface.
of progeny virions. macula An enlarged region of the inner ear containing densely
lysophospholipid Glycerolipid that contains an unesterified packed sensory hair cells specialized for detecting sound,
hydroxyl group and a single acyl chain substituent on the glycerol vibration, acceleration or gravity.
backbone. maculopapular Describes a nonvesicular skin eruption containing
lysosome A specialized membrane-bounded acidic vacuole in macules (small spots) and papules (red elevated areas of skin).
animal cells that contains hydrolytic enzymes. Material that is magnetic field The pattern of the magnetic influence derived from
internalized via endosomes is ultimately degraded in lysosomes. the presence of a magnetized entity.
lysosomotropic Describes a chemical that tends to concentrate magnetic resonance spectroscopic imaging (MRSI) A method
within the lysosomes of a cell. of acquiring nuclear magnetic resonance spectral information in a
lysozyme Antibacterial mucolytic enzyme that attacks bacterial spatially dependent manner. Data can be displayed as images in
cell walls, resulting in bacterial lysis. It is present in lysosomes and which image intensity (or colour) represents the intensity in any
is secreted in certain bodily secretions, such as tears. portion of the spectrum.
lytic Causing cell lysis. magnetosomes Intracellular membrane-bound bodies of mag-
lytic bacteriophage Bacteriophages whose infection cycle ends netic minerals present in magnetotactic bacteria.
with formation of new progeny phages and their release by cell magnetotaxis Bacterial orientation and migration along the lines
lysis. of a magnetic field.
lytic enzymes Enzymes, typically found in phagocyte lysosomes, magnitude amplification The production of a high concentration
that attack cell constituents, causing cell lysis. of active signalling molecules in response to a low concentration
lytic phage See lytic bacteriophage. of stimulus molecules.
major histocompatibility complex (MHC) A large cluster of
highly polymorphic genes that encode a family of cell-surface
glycoproteins (MHC molecules) which bind intracellularly
MA See mechanical advantage.
processed peptides and display them for recognition by T-cell
m7G cap The methylated nucleotide that is added at the 50 end of
antigen receptors. As well as controlling antigen recognition and
fully processed eukaryotic mRNA. This nucleotide, 7-methyl-
the ability to mount an immune response to a given antigen, this
guanosine diphosphate, is linked in a unique 50 –50 triphosphate
region is responsible for tissue incompatibility.
bridge with the first transcribed nucleotide in the mRNA.
major histocompatibility complex antigens See MHC mole-
m7G50 ppp50 Gm cap A modified nucleotide structure character- cules.
istic of the 50 terminus of eukaryotic mRNAs. major histocompatibility complex molecules See MHC mole-
m7GDP 7-Methylguanosine 50 -diphosphate. cules.
MAC See membrane-attack complex. malacologist Someone who studies molluscs.
Mac-1 An integrin present on leucocytes which is comprised of the malacology The study of molluscs.
subunits CD11b and CD18 and recognizes the complement malaise A general feeling of illness, body weakness and discomfort.
fragment iC3b. Also known as the complement receptor CR2. malignant melanoma A malignancy of melanocytes, almost
macrocarrier See macroprojectile. always arising in skin.
macrocyst The sexual stage of dictyostelid cellular slime moulds. management Accomplishing a defined objective by implementing
Macrocysts form from aggregations of cells in response to a plan. In the case of conservation management, a plan involving
environmental conditions of excess moisture and absence of light. a protected area or species.
Two of the cells fuse to form a giant cell that ingests all the others. mandibles The third pair of appendages on the crustacean head,
macroevolution Large-scale evolution, usually interpreted to usually reduced and specialized for food-processing.
mean all processes and patterns of phenotypic and genetic change maniraptoriform theropods Subgroup of predatory dinosaurs
at and above the species level. that includes the ornithomimids, troodontids, oviraptorids and
macrogamete Female gamete. dromaeosaurids.
macromolecular crowding The notion that a significant percen- mantle cavity The space under the shell or between the valves of a
tage of the intracellular volume is occupied by macromolecules. mollusc or brachiopod, bounded by the mantle and the anterior
macronucleus The somatic nucleus in protozoa (such as ciliates) body wall. It contains the lophophore in brachiopods.
that have nuclear dimorphism. mantle plume A body of molten material (magma) that originates in
macrophage Large phagocytic cell present in almost all tissues the Earth’s lower mantle, migrates upward through the mantle and
and derived from circulating monocytes. It scavenges tissue crust, and emerges on the surface as a geographically stable centre
debris and has effector roles in both innate and adaptive of basaltic volcanic activity (a ‘hot spot’) that can persist for millions
immunity in vertebrates. As well as phagocytosing and killing of years (e.g. the hot spot beneath the Hawaiian Island chain).
microorganisms, the macrophage can act as a professional MAP kinase Mitogen-activated protein kinase. An intracellular serine/
antigen-presenting cell to activate T lymphocytes and secretes threonine protein kinase that is part of a signalling pathway which is
numerous cytokines that influence inflammatory and immune activated by the action of extracellular signals at a variety of receptors.
responses. Granulocytes and/or plasmatocytes perform a similar it is involved in stimulating cell proliferation and gene expression.
phagocytic function in arthropods. Also known as extracellular signal-regulated kinase (ERK).
macropodid A member of the kangaroo and wallaby family of marboran The first antiviral agent against smallpox.
marsupials. marine regression A period of falling sea level.

200
Glossary

marine transgression A period of rising sea level. mechanical advantage (MA) The ratio of exerted to applied force
marker In genome mapping, any feature (such as a known DNA in a mechanical system. For example, the ratio of lever arm length
sequence) whose location in the genome can be determined. to load arm length.
marker vaccines Vaccines that allow animals that are vaccinated Meckel diverticulum A small pouch projecting from the ileum,
to be distinguished from those that have been exposed to natural first described by Johann Meckel in the eighteenth century.
infection and therefore may be carriers of the disease. meconium Fetal faeces that are dark green due to a high content of
marsupial mammals (Metatheria) The marsupials. Ovovivipar- biliverdin. The passage of meconium by the fetus in utero suggests
ous mammals, nourished from the yolk sac until hatching, when some form of hypoxic stress, which may be temporary.
the very immature young crawl out to the teats (in a pouch in Inspiration of meconium by the fetus causes airway obstruction.
many species) to complete their development. Examples are the meconium ileus Intestinal obstruction due to retained meconium.
kangaroos, opossums and bandicoots. media A layer of the pulmonary vascular wall lying between the
Marsupialia The marsupials. The name of the clade stemming endothelium and the adventitia. The media typically consists of
from the most recent common ancestor of all modern ‘pouched’ vascular smooth muscle cells.
mammals. See marsupial mammals. medial Towards, or close to, the midline.
marsupials See marsupial mammals. medical images Pictures of the human body taken using X-rays,
mass extinction Episodic events of global extent that lead to the ultrasound or any other technology in order to detect disease.
demise of numerous diverse groups of organisms in a variety of medullary cord Cellular cord between the lymphatic sinuses in
environments. They are caused by major environmental dis- the central or medullary region of a lymph node. Contains B
turbances. lymphocytes, plasma cells, reticulum cells and macrophages.
mast cell A cell that resides in tissues and mucosal surfaces and medusa The disc-like to bell-shaped sexual form (phenomorph) in
binds IgE on its surface. Binding of antigen to the cell-bound IgE the life cycle of medusozoans, commonly called a jellyfish.
triggers the release of chemicals largely responsible for the clinical megadont Large-toothed relative to body mass when compared
symptoms of allergy. with species in the same, and closely related, genera.
master sequence The dominant genomic sequence in a viral megafaunal mammals Mammalian species whose adults are
quasispecies. heavier than 40 kg.
maternal antibodies Antibodies in newborns that are passively megalopa A developmental stage of crustaceans between larva
acquired through transplacental transfer or colostral/enteral and adult.
uptake. meiofauna The aquatic fauna inhabiting the interstitial water of
maternal-effect genes Genes that are expressed by the mother soils, marine or freshwater sediments, moss cushions, etc.
during oogenesis and exert their function in the early embryo, meiosis The type of nuclear and cell division that occurs in gamete
resulting in an embryonic phenotype. formation to produce four haploid gametes (n) from a diploid
maternal-effect mutation A mutation that affects only the precursor (2n). It involves duplication of the chromosomes
progeny of a homozygous mutant mother, and is rescued by a followed by two reduction divisions, without further DNA
maternal wild-type allele. It usually represents a mutation in a synthesis, resulting in haploid cells.
maternal-effect gene. meiosis I and II Specialized cell division producing haploid germ
mating The coming together of two organisms or cells for the cells from a diploid precursor. Reduction in chromosome number
purposes of sexual reproduction. occurs during meiosis I and separation of haploid chromatids
matrix protein A structural protein of a virus particle which lies occurs at meiosis II.
under the envelope and links it to the core. meiotic drive A distortion in the transmission rate of a particular
matrix-assisted laser desorption ionization mass spectrometry chromosomal rearrangement at meiosis in favour of one allele.
(MALDI-MS) Mass spectrographic method used for rapid Certain genetic elements cause distorted segregation in hetero-
identification of proteins in which the polypeptide is applied to zygotes, such that more than 50% of gametes contain the element
a target mixed with a suitable matrix (organic acid) that generates in question.
crystals coated with protein. By laser flashes on to the target the melanocyte Specialized cell in the basal layer of the epidermis
matrix molecules are evaporated together with the peptide or which synthesizes and transfers melanin pigments to surrounding
protein molecules and yield ions that fly into a time-of-flight keratinocytes.
detector where the molecular ions are separated according to their melanoma A malignant tumour of melanocytes, usually occurring
respective masses. in the skin. Overexposure to sunlight is believed to be most
maturation (1) The final stage of differentiation of a cell or tissue, common aetiological agent.
in which it becomes fully functional. (2) The stage of viral membrane A bilayer of amphipathic phospholipids, also contain-
replication at which a virus particle becomes infectious. ing other lipids and proteins, that forms the boundary of the
maximum parsimony methods Construction of a phylogenetic cytoplasm in all living cells and also delimits intracellular
tree based on the smallest number of evolutionary changes to organelles in eukaryotic cells. The protein and, to a lesser extent,
explain the differences observed among the operational taxo- the lipid composition of biological membranes varies from cell to
nomic units. cell and compartment to compartment.
MDR Multiple drug resistance. membrane asymmetry The differences in the lipid and protein
MDR proteins Proteins that confer multiple drug resistance on composition, and physicochemical and functional properties, of
tumour cells. the two layers of a biological membrane.
measles inclusion body encephalitis An inflammatory disease membrane conductance The ability of a biological membrane to
of the brain caused by measles virus in immunocompromised conduct ions, primarily through ion channel proteins.
patients. membrane depolarization See depolarization.

201
Glossary

membrane hyperpolarization See hyperpolarization. mesenteric Pertaining to the mesentery, which is the fibrous and
membrane potential A measure of the voltage difference between vascular membrane from which the gut is suspended in the
the inside and outside of a cell membrane, which results from an abdomen.
unequal distribution of charged molecules. The membrane mesoderm The middle of the three primary germ layers formed by
potential of a cell refers to the voltage across the plasma cell gastrulation in triploblastic embryos. It lies between the ectoderm
membrane, which is usually in the range of –40 to –80 mV, and endoderm and will develop into the musculoskeletal system,
negative inside with respect to outside. circulatory system and parts of the internal organs.
membrane recycling Recovery of plasma membrane by endo- mesoglea A generally acellular layer of the body wall of
cytosis or molecular recycling. cnidarians, separating ectoderm and endoderm.
membrane resistance The ability of a biological membrane to act mesohyl That region of the sponge body enclosed by pinacoderm
as a barrier against ionic movement. The inverse of membrane and choanoderm.
conductance. mesonephros The second kidney formed in the embryo of
membrane skeleton A matrix of glycoproteins beneath the red vertebrates. It forms the adult kidney in anamniotes, such as fish
cell plasma membrane that has an essential role in maintaining and amphibia, but is superseded in adult mammals by the
the shape and flexibility of the cell. metanephros.
membrane translocator An integral membrane protein which mesoplastra Paired bony elements present in the primitive turtle
facilitates the passage of a chemical species through a membrane. plastron between the hyoplastra and hypoplastra.
membrane-attack complex (MAC) A multimeric complex of mesosome Second body section, especially that formed by the
complement components (C5b–C9) that generate a transmem- lophophore in adult phoronids and the tentacular collar in the
brane pore in eukaryotic and some prokaryotic cells, causing lysis actinotroch larva.
of the cell. messenger RNA (mRNA) An RNA molecule that represents a
membranelle A compound structure in which numerous cilia are meaningful transcript of a gene (or an operon in bacteria), that
attached to each other, forming a flattened waving structure. can be translated into protein. The mRNAs of eukaryotes encode
a single protein, whereas those of bacterial operons encode more
memory cell A B or T lymphocyte that has been activated by the
than one protein.
appropriate antigen and has not undergone differentiation into
an effector cell but has become a long-lived cell that can react messenger RNAase (mRNAase) An RNA-degrading enzyme
rapidly to re-exposure to antigen by immediately proliferating dedicated primarily or solely to degrading messenger RNAs.
and differentiating into effector cells. meta-analysis Statistical combination of the results of several
studies.
MENA A member of the VASP protein family.
metabasidium An outgrowth of a teliospore that bears basidio-
Mendelian (1) Consistent with Mendel’s laws of segregation and
spores, developing in certain basidiomycete fungi following
independent assortment of genes. (2) Applied to traits caused by a
meiosis. Also called promycelium.
single gene.
metabolic complementation The ability of the products of
Meniere disease A disorder characterized by fluctuating hearing
different defective genes to reconstitute metabolic flux of a
loss, episodic vertigo and tinnitus due to endolymphatic hydrops.
pathway. Rarely, complementation errors in the same gene give
meniscus The upward- or downward-curving boundary between a rise to intragenic complementation.
liquid and a solid surface (such as the edge of container) that
metabolic pathway A successive series of reactions catalysed by
forms as a result of capillarity.
different enzymes which result in the conversion of a molecule
menstrual migraine Migraine attacks confined to the 48 hours into one or more products with the production or consumption of
before, or the first day of, menstruation. energy.
meristems The growth regions of plants, where cells are prolifer- metabolic poisons Agents that inhibit the cell’s ability to use or
ating. The primary meristems are located at the tips of the shoot gain energy.
and root. Some cells of the meristem are perpetuated in an metabolic rate A measure of total metabolic energy use (or total
undetermined state, and thus share characteristics with animal turnover of ATP) by an organism. It is generally determined by
stem cells. measuring either oxygen consumption or metabolic heat produc-
merogony Asexual schizogony. tion.
meromictic lake A stable type of lake, independent of tempera- metabolic scope The range in rate of metabolism, expressed either
ture, built up of a mixolimnion (upper free circulating water layer) as the difference between maximal and minimal rates, or as a ratio
and a monimolimnion (lower isolated layer of water with high of maximal to minimal rates.
concentration of chemical substances) kept separate by a metabolism All of the chemical reactions within an organism that
chemocline. transform molecules and consume and generate energy.
meront Intracellular stage of sporozoan protozoan parasites, in metabolite channelling The direct transfer of a metabolite from
which asexual division usually occurs. one enzyme to another.
merosporangium (plural merosporangia) A cylindrical spor- metabotropic Describes the action of a neurotransmitter through
angium containing a row of sporangiospores. a G-protein-coupled receptor in contrast to the action of the same
merozoite Infectious stages of sporozoan parasites that are the neurotransmitter acting through ionotropic receptors.
product of merogony (schizogony). metabotropic glutamate receptors (mGluR) G-protein-
mesenchyme (1) Unpolarized cells located in connective tissue. coupled receptors for the neurotransmitter glutamate. There
(2) Non-epithelial undifferentiated embryonic cells which can are three classes of metabotropic glutamate receptors (I, II, III)
give rise to a variety of differentiated tissues. that differ in their pharmacological profiles.

202
Glossary

metabotropic receptor A G-protein-coupled neurotransmitter methanoreductosome An enzyme complex consisting of a stalk


receptor, in contrast to an ion channel-linked receptor for the and head which is attached to the inner face of the plasma
same neurotransmitter. membrane of methanogenic archaea and has methyl coenzyme M
metacentric A chromosome with a centromere at or near its reductase activity.
midpoint and two arms of about equal length. methanosomes Intracytoplasmic invaginations of the plasma
metachronal rhythm, metachronal waves Successive synchro- membrane in methanogenic archaea which have been implicated
nized waves of movement, travelling forward along the body, in methanogenesis.
typical of arthropod locomotion. methionyl-tRNA synthetase Enzyme that aminoacylates the
metacyclic Refers to a developmental form of the protozoan isoaccepting methionine tRNAs.
parasite Trypanosoma cruzi which is released in the faeces and methionyl-tRNA transformylase Enzyme that catalyses the
urine of the insect host, and is the form in which the parasite is transfer of a formyl group from N10-formyl-tetrahydrofolate to
transmitted from the invertebrate to the vertebrate host. the a-amino group of methionyl-tRNAfMet.
metagenesis See alternation of generations. methylated cytosine Cytosine in GC dinucleotides in DNA that
metal leaching Degradation of water-insoluble heavy metal salts has become modified by the addition of a methyl group.
by microbes. methylation (1) The addition of a methyl group to a molecule.
metalloenzyme An enzyme that contains a bound metal ion. (2) Modification of a cytosine residue in DNA by the addition
metalloprotease, metalloproteinase A protease that contains a of a methyl group. Methylation of some genes appears to prevent
bound metal ion. their expression.
metanephric mesenchyme Mesenchyme tissue destined to Mg branch A pathway for synthesizing chlorophylls and bacterio-
be induced by the ureteric bud to form the metanephros chlorophylls from protoporphyrin IX that branches from a
(kidney). pathway common to haem biosynthesis.
metanephridium Adult excretory organ in invertebrates, opening MHC See major histocompatibility complex.
at its inner end by one or two funnels and exteriorly by the MHC molecules Cell-surface proteins encoded by the major
nephridiopore. histocompatibility complex of genes (MHC). They bind intra-
metanephros The embryonic kidney that develops into the adult cellularly processed peptide epitopes of protein antigens and
kidney in mammals. display them on the cell surface for presentation to T cells. MHC
metaphase The phase of mitosis when the duplicated chromo- class I molecules present peptides to CD8 T cells and MHC
somes are near maximum contraction, and lie together at the class II molecules to CD4 T cells. MHC molecules themselves are
equator of the spindle. the antigens primarily responsible for the rejection of tissue
metapodials The metatarsals and metacarpals, the bones distal to grafts.
the ankle or wrist, respectively. MHC restriction Refers to the phenomenon by which an
metapopulation A set of individual populations of a species, immunocompetent T cell recognizes a foreign peptide antigen
usually within a given area, analogous to a collection of islands in only if it is presented in association with a particular MHC
an archipelago. molecule, i.e. T-cell receptors recognize unique ligands of peptide
metasome Third body section of arachnids and some crustaceans. and MHC molecule.
it is cylindrical and contains the digestive tract, gonads, MIC See minimum inhibitory concentration.
nephridia, and the main circulatory system. MIC genes MHC class I-related genes that have limited poly-
metastases Secondary tumours away from the primary site of morphism.
origin that arise as a result of the invasive spread of tumour cells. micelle (1) Small molecular aggregates formed in the presence
metastasis The spread of tumour cells through the lymph system of bile salts and allowing the absorption of lipid digestion
or blood circulation to near or distant sites where they form products into the epithelial cells. (2) A cluster of detergent
secondary tumours. molecules.
metastatic Pertaining to the transfer of disease from one part of the Michael addition Condensation of a nucleophile into an a,b-
body to another. unsaturated ketone.
metastatic tumour Secondary growth of a tumour at a site at a Michaelis complex The binary complex formed between enzyme
distance from the primary tumour. and substrate molecules preceding catalysis. It is also known as
Metatheria See marsupial mammals. the ES (enzyme–substrate) complex.
metazoa All multicellular animals whose tissues form organs. Michaelis constant Ratio between the sum rate constants of
Methanoarchaea A group of prokaryotic microorganisms, the processes leading to disintegration and to formation of a
methanogens, belonging to the domain Archaea that generate substrate–enzyme complex. It is a ‘pseudo-equilibrium’ constant
methane as a by-product of carbon fixation. related to the steady state in enzyme systems.
methanochondroitin An archaeal analogue of human chondroi- Michaelian response In an individual level of a regulatory cascade
tin, a nonsulfated polysaccharide that occurs in the cell wall of (such as an intracellular signalling pathway), an amplification
Methanosarcina. response that has Michaelis–Menten-type kinetics: e.g., as the
methanogens Prokaryotic microorganisms capable of producing stimulus increases, the resulting increment of response becomes
methane (CH4). All methanogens belong to the Kingdom progressively smaller; the response is highly graded; and it is also
Euryarchaeota of the Archaea. They subsist entirely on inorganic relatively insensitive to change.
nutrients (gaseous hydrogen, nitrogen and carbon dioxide) in the microbial fermentation Anaerobic breakdown of polysacchar-
complete absence of light and oxygen. Methane gas is released as ides and sugars by microbes to produce various products, such as
a result of the carbon fixation process. ethanol and short-chain fatty acids.

203
Glossary

microcalcification Small aggregates of calcium phosphate or midbrain The central subdivision of the brain, which contains the
oxalate occurring in secretions or in necrosis within the terminal tectum.
duct lobular unit of the glandular system of the breast, which can midgut The small intestine in many species.
produce characteristic patterns on mammography. midtarsal joint The joint in the tarsus between the talus and
microchimaerism A chimaeric condition in which an individual calcaneus (heelbone) proximally and the cuboid and navicular
contains only a small fraction of genetically different haemato- distally.
poietic and/or lymphoid cells. migraine Paroxysmal severe headache with pain-free intervals.
microdeletion The absence of a small piece of chromosome. Pulsating and throbbing qualities and unilateral distribution are
microdialysis Technique by which a substance can be measured in characteristic. Nausea and vomiting, and phobias for light and
fluid based on passage of the compound through a dialysis sound often accompany the headache.
membrane without removing actual fluid samples. migraine accompagnée See complicated migraine.
microfilament One of the three principal cytoskeletal components
migraine sine cephalgia The occasional occurrence of migrai-
in eukaryotic cells. Microfilaments are narrow (8 nm diameter)
nous auras (warning symptoms) without the accompanying
linear polymers of actin subunits. They act as architectural
headache. It usually occurs in middle-aged or elderly patients.
elements, are involved in motility, and also act as trackways for
migraine variants, migraine equivalents Unusual symptoms
intracellular transport of materials.
such as periodic abdominal pain and travel sickness, usually in
microfossils Any small fossils, such as foraminifera or spores that
children, believed to be of migrainous origin.
must be studied microscopically.
microgamete A male gamete. migraine without aura Common migraine. A throbbing, aching
microglia Resident population of bone marrow-derived cells within headache, often but not always one-sided, with sickness and
the central nervous system, resembling tissue macrophages. photophobia, but no specific warning symptoms (aura).
microinvasion Foci of invasive carcinomas, none which exceed migrant pool colonization The founding of a new population
1 mm in diameter, arising from ductal carcinoma in situ. with colonists chosen at random from all populations in the
micrometeorite A particle from space that is small enough to be metapopulation.
slowed down when it reaches the Earth’s atmosphere without migration In the context of population genetics, this term refers to
being burnt up. gene flow and is measured as that proportion of a local
micronucleus The generative (sexual) nucleus in protozoa, such as population that arrives in each generation from other popula-
ciliates, that have nuclear dimorphism. tions of the same species.
micronutrient An essential mineral nutrient that is only required Milankovitch cycles Astronomically determined climatic cycles
in small quantities. Tissue concentrations of micronutrients are with dominant periodicities of 100 000, 41 000 and 22 000 years.
equal to or less than 100 mg per kg dry matter (or less than 0.005% Milankovitch theory Explanation of the timing of glacial–inter-
body weight). glacial climate based on changes in the Earth’s orbit.
microorganisms Living organisms that are too small to be seen Miller indices In X-ray crystallography, a set of three integer
with the naked eye, e.g. bacteria, fungi, protozoa and single-celled values, denoted (h,k,l), used to label a reflection or structure
algae. Viruses are also considered as microorganisms. factor according to the direction and angle of scattering of the
microphyllous leaves Elongate leaves with a single (or occasion- X-ray beam.
ally two parallel) veins running along their length. miniature postsynaptic potential (MPSP) A postsynaptic
microprojectile/microcarrier Any microscopic and coherent par- potential caused by the spontaneous secretion of a single
ticle that when sufficiently accelerated can penetrate cells and tissues. presynaptic vesicle of transmitter.
micropylar end The part of a seed in which the embryo grows. minimal effective concentration An in vitro measure that defines
micropyle The small channel through which a pollen tube enters an the effective concentration of an agent against a fungal organism.
ovule prior to fertilization. minimal medium The standard medium having the fewest and
microsatellites Tandem arrays of very short, moderately simplest compounds upon which an organism will grow well.
repetitive, localized DNA sequences (e.g. (CA)15) present in
minimal inhibitory concentration See minimum inhibitory
mammalian and other genomes.
concentration
microscope slide A rectangular sheet of thin glass used to support
minimally invasive therapy The performance of an interventional
thin sections or smears of the specimen for optical microscopy.
procedure with the minimum of trauma to the patient. Also
microspike See filopodia.
known as minimal access surgery.
microstromatolite Fossilized layers of bacterial mats and sedi-
ment, producing a very finely laminated rock stratum. minimum resolved distance (MRD) The least distance between
microtomy Use of a mechanical device (a microtome) to cut thin two points that can be resolved by an imaging system.
sections for microscopy from a larger specimen. minimum viable population The minimum size of population
microtubule-organizing centre (MTOC) The major origin of that ensures a greater than 95% chance that the population will
microtubules in a living cell. persist for 100 years into the future.
microtubules One of the three principal cytoskeletal components minisatellites Tandem arrays of moderately repetitive, localized
of the eukaryotic cell. Microtubules are stiff hollow tubes of sequences with repeat size ~15 bp, longer than that of
protein (25 nm in diameter) composed of tubulin subunits. microsatellites.
They act as architectural elements and also as trackways for minus end The end of a microtubule that is slowest growing.
intracellular transport of materials. Movement of cilia and MIP Major intrinsic protein of bovine lens. It was the first defined
flagella is based on microtubules, and they also form the mitotic member of a family of presumed channel proteins that includes
and meiotic spindles. the aquaporins.

204
Glossary

missense error An error in translation that results from a codon modulus A measure of the stiffness of a material, which helps
being incorrectly decoded. It results in the incorporation of the determine the degree to which it deforms if forces are applied.
incorrect amino acid into the protein. molecular chaperones Proteins that aid in protein folding,
missing self model The hypothesis that natural killer cells survey protein associations or protein transport, but are not part of
cells for the presence of MHC class I molecules, killing those in the final protein complex.
which it is missing. Originally proposed in 1985 and now molecular clock The theory that mutations accumulate in DNA
supported by a large body of evidence. sequences at an approximately constant rate, and thus can be
mitochondria (singular mitochondrion) Energy-generating used as a measure of the absolute time of divergence of two
organelles of eukaryotic cells. They are bounded by two sequences. It it most likely to hold for sequences that still encode
membranes, the inner one being folded into cristae that carry proteins with the same function.
the electron transport chains that generate the energy for the molecular marker See marker.
production of ATP by oxidative phosphorylation. The tricar- molecular mimicry (1) The case when a parasite antigen resembles
boxylic acid cycle occurs in the central mitochondrial matrix. that of a host antigen. In some cases this may mean that an
Mitochondria carry a small DNA genome that encodes some immune response is not mounted against that parasite. In other
mitochondrial proteins. cases, it can result in a transient autoimmune reaction as
mitochondrial bottleneck Genetic term to explain how a antibodies evoked by the parasite cross-react with the host
mitochondrial DNA polymorphism can rapidly become pre- antigen. (2) Similarity in the three-dimensional structures of
dominant between generations. It is suggested that only a small divergent molecules which facilitates recognition by common
subset of mitochondrial DNA molecules is selectively amplified elements.
during oocyte maturation. molecular phylogenetics The study of the evolution of molecules
mitochondrial DNA (mtDNA) Small circular DNA molecules (DNA and proteins), generally by constructing phylogenetic trees
present in mitochondria, and which encode some mitochondrial from DNA and amino-acid sequences.
proteins. molecularity The number of molecules of reactants involved in the
mitochondrion See mitochondria. transition state of an elementary chemical process.
mitogen A chemical or physical agent that nonspecifically induces monoamines The neurotransmitters dopamine, noradrenaline
cells to enter mitosis. (norepinephrine), serotonin and adrenaline (epinephrine).
monobrachial homology Sharing of a chromosome arm in
mitogen-activated protein kinase See MAP kinase.
double Robertsonian heterozygotes.
mitoplast A mitochondrion with a ruptured outer membrane that
monociliated epidermis Epidermis in which each epidermal cell
has lost the soluble content of the intermembrane space but is
has only one cilium.
otherwise intact.
monocistronic Describes a messenger RNA that encodes a single
mitosis A type of nuclear division that produces two daughter
protein.
nuclei containing chromosome sets identical to each other and to
monoclonal antibody Antibody of one specificity produced by a
that of the parent cell.
single clone of antibody-secreting cells. They can be produced in
mitotic parthenogenesis Development of an individual from an culture by fusing a single B cell with a myeloma cell to yield a
unfertilized egg not undergoing meiotic divisions. Such indivi- specific antibody-secreting hybridoma cell line.
duals are genotypically identical to the mother.
monoclonal gammopathy Pertaining to or originating in a single
mixed lymphocyte reaction A test for histocompatibility. When clone of B cells. All the cells in such a clone will produce an
lymphocytes (effectively T cells) from two incompatible people identical specific immunoglobulin.
are cultured together, they will recognize each other as foreign monoclonal (M) component Immunoglobulins produced by a
and start to proliferate and differentiate into helper T cells and plasma cell dyscrasia. These are seen as a sharp discrete band on
cytotoxic T cells. This test mimics allograft recognition. protein electrophoresis.
mixed semidiurnal tides A sequence of tides consisting of two monoclonal Arising from a single clone of cells.
high and two low tides over approximately 24 hours and 50 monocotyledonous Belonging to the group of plants called
minutes. The high tides are of unequal amplitude, as are the low monocotyledons, which have a single cotyledon in the seed.
tides. monocotyledons A major division of the flowering plants having
mixotrophic Capable of using two or more modes of nutrition (e.g. only one seed leaf or cotyledon. Most monocotyledons have
phototrophic and heterotrophic) during the life cycle, as in some narrow, parallel-veined leaves, as in grasses and lilies.
protists. monocyclic cascade A simple system where a signalling protein A
mixotrophy Nutritional mode exhibited by some protozoa and can be converted between active and inactive forms by opposing
other protists in which two or more methods of satisfying activator and inactivator enzymes.
metabolic needs, e.g. photoautotrophy and heterotrophy, may be monocytes Circulating white blood cells that are the source of
combined in a single organism. tissue macrophages.
mmc Megaspore mother cell, a diploid cell that gives rise via monocytoid B cells B cells with the cytological appearance of
meiosis to four megaspores. histiocytes, which may collect in lymph node sinuses in some
mobile phase Buffer used to equilibrate affinity chromatography reactive states.
columns. monodisperse suspension of particles A suspension of particles
mobilization Conjugal transfer of a plasmid lacking a complete set of very uniform diameter. They are all the same size and therefore
of transfer functions, which is not capable of autonomous alter in an exactly reproducible fashion the properties of cells to
transfer and requires transfer functions supplied by a coresident which they attach, e.g. the magnetic properties if they themselves
conjugative plasmid. are magnetic.

205
Glossary

monogenic Applied to traits that are caused by a single gene. morula The mammalian conceptus at the 16–32-cell stage.
monomer (1) A single subunit of a dimer or a larger multimeric mosaicism Condition in which an individual is composed of cells of
complex. (2) A single repeating unit of a polymer. two different genotypes.
mononegavirales Order of viruses consisting of four families with mosasaurs A group of large, extinct marine monitor lizards that
nonsegmented negative-stranded RNA genomes. might be closely related to snakes.
mononuclear With a single nucleus. Applied, e.g. to those white motif A characteristic or conserved amino-acid sequence contain-
blood cells, such as monocytes, that have a single round nucleus. ing functionally important residues.
mononuclear phagocyte system A family of widely distributed motional freedom The ability to change conformation. In an
macrophage-related cells, usually phagocytic, that are derived amphipathic lipid, the hydrocarbon chains have considerable
from blood mononuclear cells, as opposed to polymorphonuclear motional freedom at their methyl termini and less towards the
(neutrophil) leucocytes. This term has not entirely replaced the polar headgroup.
older term, reticuloendothelial system. motional order Lipid hydrocarbon chains have considerable
monophyletic Describes a phylogenetic group (a clade) that conformational freedom at their methyl termini, and the
consists of a common ancestor and all of its descendants. motional order is low. Conversely, their degree of conformational
Accordingly, such a group should be definable by the possession freedom towards the polar headgroup is low and the motional
of one or more shared characters derived from the ancestor order there is high.
(synapomorphies). motor learning The adaptive changes occurring in the brain that
monophyletic group A group of species that includes their single allow movements to be executed skilfully, and to be well adjusted
common ancestor and all of this ancestor’s descendants. to the mechanical loads that must be moved.
monophyletic origin Describes a trait that has been established motor proteins Enzymes that move along the surface of micro-
once and then transmitted to different systematic categories tubules or microfilaments. Depending on the other structures
during evolution. with which they interact, they can move cytoplasmic constituents
monophyly See monophyletic. along the cytoskeletal polymers, or they can move the cyto-
monosaccharide A molecule consisting of a single sugar. skeletal structures themselves.
monosome A messenger RNA molecule with a single attached moult cycle The time between two moults (ecdyses). Divided into a
ribosome. period of preparation for moult (called pro-ecdysis) and a period
monosomy A condition in which one chromosome of a pair is of normal life activities (called anecdysis).
missing. moulting In arthropods, the process of shedding the cuticle at
monoxenic culture A culture in which only one species of bacteria periodic intervals during the growth period.
or other food organism is allowed to be present with a cultured mounting Immersion of a stained tissue section or smear,
protist. attached to a microscope slide, in a medium of high refractive
monoxenous life cycle The life cycle in which the sexual and index, the whole usually being overlaid with a thin glass
asexual stages of a parasite occur in the same host. coverslip.
monozygotic twins Genetically identical twins, derived from the
Mr Relative molecular mass.
same egg.
MRD See minimum resolved distance.
morph A phenotypic variant, usually one of two or more possible
mRNA See messenger RNA.
developmental outcomes.
mRNAase See messenger RNAase.
morphallaxis Regeneration process by which new structures are
formed in the absence of cell proliferation. It relies on migration mRNA stability determinant A segment of an mRNA that helps
and terminal differentiation of precursor cells or possibly on to determine the mRNA half-life.
transdifferentiation. mRNA turnover The rate at which an mRNA is degraded, usually
morphogen Molecule that is distributed in a gradient and defined in terms of mRNA half-life.
subdivides a field of cells into more than two domains. MRS See nuclear magnetic resonance spectroscopy.
morphogenesis The development of form, or morphology, in MSP Major sperm protein, a protein from the sperm of certain
organisms during embryogenesis. nematodes (e.g. Ascaris) that polymerizes to drive the cell forward
morphogenetic movement Cell or tissue migrations that lead to in a manner similar to that proposed for actin in most other
changes in organ or body shape. eukaryotic cells.
morphogenetic phenomena Developmental processes asso- mtDNA See mitochondrial DNA.
ciated with cell division in ciliates. These include the development MTOC See microtubule-organizing centre.
of a new oral apparatus (stomatogenesis), assembly and mucopolysaccharides Gelatinous carbohydrates of high Mr
positioning of new organelles, as well as degradative processes. (~5  106) that both lubricate and serve as a sticky cement.
morphological species concept The differentiation of species on mucormycosis Invasive infection caused by the fungus Mucor
morphological grounds, providing the baseline for taxonomic racemosus.
description. mucosa (plural mucosae) General term for the secretory epithelia
morphology (1) The shape and form of a cell or organism. (2) The lining the gastrointestinal, respiratory and urinogenital tracts.
scientific discipline that studies the diversity of shapes of multifunctional protein A protein containing two or more
biological organisms. discrete and different enzyme catalytic centres, on separate
morphometrics The application of multivariate methods to the domains.
analysis of size and shape in organisms. multigene family Family of DNA or amino-acid sequences in
morphotype A taxon diagnosed by a unique combination of which some sequences or groups of sequences have evolved by
characters. gene duplication events.

206
Glossary

multinet growth hypothesis Physical explanation that plant cell mycoheterotroph A plant that in the juvenile stages or through-
elongation results from passive reorientation of the cellulose out its life lacks chlorophyll (i.e. is heterotrophic). Instead of
microfibrils from transverse to longitudinal in the outer layers of photosynthesizing it receives all of its carbon by way of its
the wall to affect growth along the entire axis. mycorrhizal fungal symbionts.
multiphoton microscope A microscope in which the illumination mycoparasite A fungus that parasitizes another fungus.
is confined to a spot in the specimen by means of two- or three- mycorrhiza (plural mycorrhizae) A symbiotic relationship
photon excitation of a fluorescent probe using a high-power between a fungus and plant roots.
pulsed laser. mycorrhizal mushroom Describes a mushroom-forming basidio-
multiple alleles The existence in a population of more than two mycete fungus, whose mycelium forms a mutually beneficial
variant forms of a gene at a given locus. association with the roots of a plant. Thus, the mushroom is
multiple cloning site A short sequence of DNA that contains the typically found near the plant host.
recognition sites for several restriction enzymes and thus mycorrhizosphere The zone surrounding the mycelium of a
can accept DNA fragments cut with a variety of restriction mycorrhizal fungus as it extends into soil.
enzymes. mycosis (plural mycoses) Any disease caused by a fungus.
multiplicity of infection Number of infectious particles per host mycotroph A plant that is dependent upon, or highly responsive
cell in the initial infective dose. to, the provision of nutrients by a fungus.
multipotential progenitor A cell that can produce daughter cells myelin A discontinuous fatty insulating sheath formed around the
that will give rise to multiple lineages. axons of certain neurons, which facilitates fast nerve conduction.
multiregional hypothesis The hypothesis that modern humans It is produced by Schwann cells in the peripheral nervous system
are diffusely descended from precursor populations, called and oligodendrocytes in the central nervous system, which wrap
archaic Homo sapiens, distributed over much of the tropical their cell processes around the axon, forming a sheath of layers of
and temperate Old World. membrane.
multispot array A set of immobilized capture molecules (probes) myeloid progenitor A cell that is committed to a pathway that will
that participate in a solid–liquid interfacial interaction with target give rise to mononuclear and polymorphonuclear leucocytes.
molecules contained in the sample solution applied to the surface myoblasts Undifferentiated cells that develop into muscle cells
of the multispot array. during development.
multituberculate mammals An extinct order of mammals, myocarditis Inflammation of the myocardium or the muscle of the
common in the Mesozoic era, that were ecologically equivalent heart.
to modern-day rodents. myoclonus Pathological sudden involuntary muscle jerking.
murein Cell wall peptidoglycan of the Bacteria. myofibroblast A fibre-forming cell of the connective tissue, which
mutagenesis The process of causing a change in a DNA contains large amounts of contractile assemblies of smooth
sequence. muscle actin and myosin. Myofibroblasts are regarded as
mutagens Agents that will induce a change in a cell’s DNA. contractile cells.
mutant spectrum The set of variant genomes present in an myofilaments Actin and myosin protein filaments that are linked
evolving quasispecies. by cross-bridges and slide relative to each other to produce force.
mutation A permanent heritable change in the genetic material of a myoglobin A haemoprotein that reversibly binds dioxygen and
cell or organism. This could be a change in the base sequence of stores it in muscle tissue.
the DNA that affects a single gene, or a change in the number or myopathy A neurological disorder affecting muscles.
structure of the chromosomes. In classical genetics, the term myosin Any of a superfamily of motor proteins (17 classes
usually refers to a change that has a demonstrable effect on the identified so far) present in probably all eukaryotic cells. They
phenotype. bind to actin and use the energy of ATP hydrolysis to generate
mutualism Persistent and intimate association between two (or force and movement along actin filaments.
more) dissimilar organisms that provides advantages, particu- myotube Multinucleate cell that develops from fusion of several
larly in terms of fitness, to all partners. myoblasts and develops into a skeletal muscle cell.
mutualistic interaction An interaction between organisms in myotubules The earliest muscle structures formed during devel-
which the darwinian fitness of both (or all) interacting species is opment.
increased as a result of the interaction. myristoylation The covalent attachment of a myristoyl group
mutualistic symbiosis A symbiosis that is beneficial to all the (derived from the fatty acid myristic acid) to the N-terminal glycine
partners in the association. residue of a nascent polypeptide. This posttranslational modifica-
myalgia Muscle pain. tion can provide a way of attaching a protein to a membrane.
Myb proteins, MYB proteins Transcription factors structurally myrmecophagous Feeding on ants and termites.
homologous to the mammalian cellular proto-oncogene c-Myb.
They have common DNA-binding domains that are involved
in the control of gene expression of various metabolic and NA See numerical aperture.
developmental processes. Na1 , K1-ATPase A transmembrane protein with ATPase activity
Myb-related transcription factor See Myb proteins. that uses the energy released by hydrolysis of ATP to transport
mycelium The vegetative part of a fungus, composed of a network three Na1 out of the cell and two K1 into the cell per ATP
of hyphae. hydrolysed. It is chiefly responsible for generating the membrane
mycetocyte Also called mycetome, an evagination of the midgut potential in animal cells, maintaining the ion gradients that are
of certain insects. used by other transport proteins.

207
Glossary

NADP1 Oxidized form of nicotinamide adenine dinucleotide negative-stranded RNA virus A virus whose RNA genome is
phosphate. complementary to its mRNA.
NADPH Reduced form of nicotinamide adenine dinucleotide neighbour-joining analysis A statistical method that sequentially
phosphate, a small water-soluble molecule that acts as a hydrogen identifies neighbour pairs (operational taxonomic units con-
carrier in biochemical reactions. nected through a single internal node) that minimize the total
NADPH oxidase A multicomponent enzyme that oxidizes intra- length of the phylogenetic tree.
cellular NADPH (the reduced form of nicotinamide adenine nektic, nektonic A term applied to aquatic organisms that actively
dinucleotide phosphate) to NADP+ and reduces molecular swim.
oxygen, O2, to superoxide anions, O 2. nematocyst Any of 30 types of adhesive, ensnaring, coiling and
naive lymphocyte Lymphocyte that has not previously been penetrating cnidocysts present in cnidarians.
stimulated by its specific antigen. nematodes A large phylum of organisms commonly known as
nascent polymer A polymer molecule in the process of being round worms. They include the model organism Caenorhabditis
synthesized. elegans.
nascent strand A polymer strand in the process of being Neognathae A taxonomic group that includes all modern extant
synthesized. birds. Members of this group have a prokinetic skull, keratin-
native Describes the three-dimensional conformation of a protein covered and toothless jaws (the beak) and several other characters
or peptide that represents the biologically active structure. such as a keel on the sternum (breastbone) for the attachment of
natriuretic Causing an enhanced tendency for urinary excretion of large flight muscles. These characters separate this taxon from the
sodium. Palaeognathae.
natural cytotoxicity The ability of natural killer (NK) cells to kill neonatal alloimmune thrombocytopenia purpura Destruction
certain target cells without preactivation. Also known as of platelets of the fetus or the newborn by platelet-specific
spontaneous cytotoxicity. alloantibodies derived from the mother.
natural immunity See innate immunity. neonate Newborn. In humans, it refers to an infant of up to 4
natural killer cell (NK cell) A type of antigen-nonspecific non-T, weeks old.
non-B cytotoxic lymphocyte that plays a crucial role in innate neoplasia Abnormal proliferation of cells (‘new growth’). The
immunity to viruses and other intracellular pathogens. Unlike word generally refers to tumour cell growth and behaviour that is
cytotoxic T lymphocytes, NK cells can lyse target cells without initiated by genetic alteration, and which persists after removal of
the need for previous antigen recognition or sensitization. NK the agent that caused the change.
cells are also able to kill certain tumour cells, and are the cells neoplasm An abnormal growth, caused by proliferation of cells.
mediating antibody-dependent cell-mediated cytotoxicity. The term is synonymous with tumour, either benign or
natural killer cell receptor (NK cell receptor) Receptors on NK malignant.
cells that can either activate or inhibit the cytotoxic activity of the neoteny (1) The persistence of embryonic characters into adult-
cell. Examples are KIR (killer cell inhibitory receptor) and KAR hood. (2) The attainment of sexual maturity in an immature stage.
(killer cell activating receptor). nephridium (plural nephridia) An invertebrate organ that serves
natural product A compound produced by a living organism. excretory or osmoregulatory functions.
natural selection The evolutionary process by which the organ- nephron An individual functional unit of the vertebrate kidney.
isms in a population that are best adapted to the environment It is composed of the glomerulus and a tubule draining the
increase in frequency relative to less well-adapted forms over a glomerular filtrate.
number of generations. nephrotoxicity The property of being toxic to kidney cells.
naturalized Describes introduced (alien) species that have formed nerve regeneration Regrowth of nerve fibres after injury.
self-sustaining populations in the wild at their new location. nerve ring A visible ring of fibrous material of the nervous system
nauplius larva The characteristic larvae of crustaceans, with a which usually encircles the nematode pharynx.
median simple eye and three pairs of appendages, which will NES See nuclear export signal.
become the antennae and mandibles of the adult. network A system of wires and communication standards that
NBS See nucleotide-binding site. allow computers to communicate with one another.
necrosis Pathological cell death and tissue destruction, usually neural activity The all-or-none electrical discharge of a neuron
in a localized area, which is due to cell lysis as opposed to that is propagated along its axon with the resultant release of
programmed cell death due to apoptosis. neurotransmitter from synaptic vesicles.
necrotizing Causing cell or tissue death. neural crest An embryonic tissue that gives rise to many tissues,
needle core biopsy A biopsy taken by inserting a sharpened including the neurons and glia of the peripheral nervous system.
cutting needle into a tissue. neural plate The flattened plate of neural ectoderm on the dorsal
negative chemotactic stimulation A decreasing gradient of side of a vertebrate embryo that will give rise to the nervous
chemoattractant or an increasing gradient of chemorepellent. system. It becomes evident after gastrulation and neural
negative eugenics Use of programmes, techniques or social induction.
policies with the goal of reducing or eliminating negative (harmful neural tube In vertebrate embryos, the precursor to the central
or undesirable) genes in a population, often by preventing nervous system which is formed when the lateral edges of the
reproduction of individuals believed to be carrying such genes. neural plate roll dorsally and fuse at the dorsal midline to form a
negative selection The process whereby self-reactive clones of T tube.
cells or B cells are deleted from the developing lymphocyte neuraminidase An enzyme with sialidase activity. It is present in
repertoire, principally by apoptosis. some viruses.

208
Glossary

neuroblast An embryonic progenitor cell of a neuron. In Newtonian flow Flow of a fluid whose viscosity does not change
arthropods, neuroblasts can also give rise to glial cells as well as with changes in shear rate or over time.
to neurons. In vertebrates, the name refers to the postmitotic but Newton’s rings A circular pattern of fringes with alternating
immature neuron. intensity that arises from the interference of monochromatic light
neurofilaments Intermediate filaments of neurons, which act as partially reflected from a flat glass surface with the light partially
internal support for the axon. reflected from a concave glass surface.
neurogenesis The ‘birth’ or formation of new neurons during NF-kB Nuclear factor kB. A transcription factor that enhances
embryonic development. It occurs in the ventricular zones lining transcription of a number of genes, particularly those involved in
the lumen of the neural tube. inflammation and immunity.
neurogenic Describes genes whose loss of activity (defined niche specialization The particular habitat and mode of life
originally by mutants) leads to neural hypertrophy. sustenance of a given species.
neuromast Individual sensory organs of the lateral line, composed niche The geographic location, habitat, activity pattern and diet
of hair cells and supporting cells. that characterizes a given species (or group of species) and
neuromodulator A neurotransmitter whose main effect is to according and to which it is particularly adapted. See also
modify the properties of an ion channel that primarily responds fundamental niche, realized niche.
to another factor, such as voltage. nick A discontinuity in the phosphodiester backbone of duplex
neuron A nerve cell, the principal information-processing and DNA in which no base pairs are missing.
signalling cell of the nervous system. Neurons develop elongated nicotine An alkaloid that acts as a nonphysiological agonist at the
processes, dendrites and axon, that receive and transmit ion channel type of acetylcholine receptor, which is thus known as
information, respectively. Information travels as electrical signals the nicotinic acetylcholine receptor (nAChR).
within neurons and is transmitted between neurons by chemical Nieuwkoop centre A region of the dorsal endodermal mass in the
neurotransmitters released at specialized cell junctions called Xenopus blastula that induces the adjacent marginal zone cells to
synapses. become dorsal mesoderm (notochord and paraxial mesoderm).
neuropathy A neurological disorder affecting the peripheral nif Gene designation for the molybdenum nitrogenase system.
nerves. nitric oxide A short-lived membrane-permeable signalling mole-
neuropil Regions of densely intertwined dendrites and axons of cule.
neurons, excluding the cell bodies. nitrogen fixation The conversion of molecular nitrogen (N2) into
neuroscience The study of the brain and nervous system. metabolically usable nitrogen, principally NH+ 4 . Biological
neurotoxicity The disturbance of neuronal function or causation nitrogen fixation can be carried out only by a few species of
of neuronal death by an external agent. free-living and symbiotic bacteria.
neurotransmitter A signalling molecule released from the pre- NK cell See natural killer cell.
synaptic axon at a synapse after the arrival of an action potential NK cell receptor See natural killer cell receptor.
at the axon terminal. It diffuses across the synpatic cleft and, NLS See nuclear localization signal.
depending on the type of neurotransmitter and neuron involved, NMDA receptor A type of receptor that responds to the excitatory
can cause either depolarization or hyperpolarization of the neurotransmitter glutamate and which is distinguished from
postsynaptic cell. other glutamate receptors by its selective response to N-methyl-
neurotrophins Proteins important for neuronal growth and D-aspartate and activation of calcium currents.
survival. An example is neuronal growth factor (NGF). They NMDA N-methyl-D-aspartate. A chemical used to distinguish a
act at specific receptors (neurotrophin receptors). particular type of glutamate receptors.
neurulation Formation of the neural tube during embryogenesis NMR See nuclear magnetic resonance.
by the rolling up of the neural plate and fusion of the neural folds. NMRS See nuclear magnetic resonance spectroscopy.
neutral allele, neutral mutation A mutation that has no NNS RNA virus Nonsegmented negative-strand RNA virus
apparent favourable or unfavourable effect on phenotype and nomenclature The assignment of taxonomic rank to the groups
whose fate in a population is determined solely by random genetic resulting from classification and naming the organisms according
drift. to their rank.
neutral substitutions Base changes in the reading frame of a gene nominalism The philosophical thesis that individuals are real but
that do not alter the encoded amino-acid sequence. classes are not real, sometimes associated with the notion that all
neutral theory The proposal that the majority of sequence groups are purely arbitrary or mental constructs.
differences observed between the DNA or proteins of two species noncanonical splice site Unconventional splice site.
are the result of fixation of neutral or nearly neutral mutations by noncentromeric chromosome A DNA molecule (with asso-
random genetic drift, rather than the result of natural selection. ciated protein) that lacks spindle attachment regions (centro-
neutropenia A reduction in the blood neutrophil count below meres). Such entities are sometimes considered not to be true
normal. chromosomes, and then are called ‘autonomously replicating
neutrophil A phagocytic white blood cell with a distinctive lobed pieces’.
nucleus and granules that stain with neutral dyes. It is one of the nondisjunction Failure of chromosome pairs or sister chromatids
first defences against many types of bacterial infection, ingesting to separate at cell division, resulting in aneuploidy in the daughter
and killing bacteria. Acting in both innate and adaptive cells.
immunity, neutrophils bear complement and Fc receptors and nonhomologous end-joining enzyme One of a group of
migrate from the blood into sites of infection or inflammation. enzymes that form complexes capable of joining, or religating,
neutrophiles Microorganisms that grow with a pH optimum near double-stranded ends of DNA which has been cleaved or broken,
neutrality. when the double-stranded ends have no sequence homology.

209
Glossary

nonhomologous recombination Recombination between DNA nuclear localization signal (NLS) Sequence of amino acids that
molecules sharing little or no homology. Also known as end- targets a protein for import into the nucleus.
joining. nuclear localization Present in the cell nucleus.
nonpersistent transmission See stylet-borne transmission. nuclear magnetic resonance (NMR) A technique that examines
nonselective cation channel An ion channel that is selective for the structure of molecules by detecting nuclear spin reorientation
cations (positively charged ions) over anions (negatively charged (radiofrequency transitions) in an applied magnetic field.
ions), but shows little selectivity between Ca21 , Na1 and K1 . nuclear magnetic resonance spectroscopy (MRS/NMRS) A
nonself Everything that is not part of the body’s own tissues and branch of spectroscopy operating in the radiofrequency region of
may be recognized by the immune system. the electromagnetic spectrum, roughly from 0 to 109 Hz. The
nonsense codons Codons that do not specify an amino acid but signal arises from the interaction between atomic nuclei and a
signal the end of the region of an mRNA to be decoded. Also magnetic field.
known as termination or stop codons. nuclear pore complex (NPC) A large multiprotein complex
nonsense suppressors tRNAs with anticodon mutations that inserted into the double membrane around the nucleus that
allow translation of stop codons. mediates all communication between the cytoplasm and the
nonsense-mediated mRNA decay The process whereby the cell nuclear interior.
recognizes and rapidly degrades mRNAs with premature nuclear receptors Receptor proteins, such as those for some
nonsense (termination) codons. steroid hormones, that are located within the nucleus, or
nonstructural proteins Virus-encoded proteins that function in translocate to the nucleus after binding their ligand.
the replication of the virus and are not part of the viral particle. nuclear reprogramming The resetting of the pattern of gene
nonunion distal (NUd) A type of deletion in sister chromatids in expression that occurs in a differentiated somatic cell nucleus
which the broken ends of the remaining sister chromatids fuse but following its transfer to an enucleated oocyte. After reprogram-
the broken ends of the deleted acentric portions do not. ming the nucleus can support normal embryogenesis.
nonunion proximal (NUp) A type of deletion in sister chromatids nuclear transfer The method of reconstructing a one-cell
in which the broken ends of the remaining sister chromatids do embryo from an enucleated oocyte and the nucleus of a
not fuse, but the broken ends of the deleted acentric portions do. donor cell to generate an embryo and ultimately a cloned
nonzonal bone Bone tissue that does not show zonal ‘growth animal.
rings’. nuclease An enzyme that degrades nucleic acids (DNA or RNA)
noradrenaline The catecholamine neurotransmitter of the sympa- by cleaving phosphodiester bonds. Endonucleases attack internal
thetic nervous system. Called norepinephrine in North America. bonds of the polymer, whereas exonucleases attack from the end,
norepinephrine See noradrenaline. usually liberating mononucleotides.
NOR See nucleolar organizing region. nucleocapsid The nucleoprotein core of a virus, consisting of the
norm of reaction See reaction norm. nucleic acid genome and associated proteins.
normal pressure hydrocephalus (NPH) Syndrome of ventricular nucleoid In prokaryotes, the region of the cytoplasm occupied by
enlargement with normal intracranial pressure, characterized by the DNA and associated proteins.
a symptom complex of dementia, gait instability and incon- nucleolar organizing region (NOR) The chromosomal regions
tinence. that form the nucleolus in interphase eukaryotic cells.
nosocomial infection Infection pertaining to or originating in nucleolus Region in the nucleus where rRNA is transcribed and
hospital. ribosomes partially assembled.
notochord A rod-like mesodermal structure that forms just ventral nucleophile An atom (or chemical group) with an available lone
to the neural plate and tube in vertebrate embryos. It provides a pair of electrons that can act as an electron donor in a chemical
primitive backbone for the embryo. reaction. Common active-site nucleophiles in enzymes are the
novelty A new structural character arising in a phylogenetic hydroxyl group of serine and threonine, the sulfhydryl group of
lineage that is neither homologous to any structure in the cysteine, and the imidazole group of histidine.
ancestral species, nor serially homologous to any other structure nucleophilic attack The reaction of an electron-rich atom
within the same organism. with an electron-deficient atom to form a bond. An example
NOx The nitrogen oxides (NO, NO2, N2O, N2O3, etc.) collectively. is the attack of the e-amino group nitrogen of a lysine residue
In literature on the effects of air pollutants on vegetation, often of DNA ligase on the a-phosphorus of ATP to give a P–N
only NO and NO2 are referred to as NOx as these two substances bond.
contribute most to the nitrogen load of terrestrial ecosystems. nucleophilic Describes an electron-rich atom or chemical group
NPC See nuclear pore complex. that readily reacts with compounds (electrophiles) containing
NPH See normal pressure hydrocephalus. positive charge or that are electron deficient.
nt An abbreviation for nucleotide(s). nucleophilic substitution Chemical reaction whereby a nucleo-
nuclear DNA The DNA found in the chromosomes contained in phile attacks an atom and often leads to the dissociation of
the nucleus of a eukaryotic cell, as opposed to the DNA of another group covalently bound to that atom, called the leaving
mitochondria and chloroplasts. group.
nuclear export signal (NES) Sequence of amino acids that targets nucleoporin A protein component of the nuclear pore complex.
a protein for export from the nucleus. nucleoprotein filament A complex of RecA protein, DNA and a
nuclear lamina A layer of proteins (the lamins) on the inner side of nucleoside triphosphate cofactor.
the nuclear membrane, to which nuclear pore complexes are nucleoside Molecule consisting of a purine or pyrimidine base
thought to be anchored. attached to a pentose sugar (e.g. ribose, deoxyribose).

210
Glossary

nucleosome The basic structural unit of eukaryotic chromatin, Ohm’s law The relationship between current (I), voltage (V) and
consisting of an octamer of histone molecules ((H2A, H2B)2 resistance (R): a current of 1 ampere (1 coulomb s1) flowing
(H32, H42)) plus one molecule of H1, around which is wrapped through a resistance of 1 ohm will generate a voltage drop across
about 1.7 turns (180–200 base pairs) of DNA double helix. that resistor equal to 1 volt (V 5 IR).
nucleosome core particle The histone octamer and the length of Okazaki fragment A short length of DNA that is synthesized
DNA (146 base pairs) wrapped round it that resists digestion by during discontinuous synthesis on the lagging strand of replicat-
micrococcal nuclease. ing DNA. Okazaki fragments are 1000–2000 nucleotides in
nucleosome position Nonrandom contacts of histones with DNA prokaryotes and 50–200 nucleotides in eukaryotes. After their 50 -
with respect to sequence. RNA primers are removed, the fragments are joined together by
nucleotide excision repair A DNA repair process whereby DNA ligase to form a complete DNA strand.
damaged nucleotides in the cell’s DNA are removed as part of a Oldowan The name of a type of stone tool industry that comprises
short single-stranded DNA fragment and the integrity of the crude chopping tools and flakes.
DNA restored to its normal state by new DNA synthesis using the oligoastrocytoma A brain tumour arising either from an oligo-
exposed normal strand as template. dendrocyte or an astrocyte.
nucleotide-binding site (NBS) Amino-acid residues that form oligoclonal Made up of only a few clones, as, e.g., an immune
the binding pocket for nucleoside triphosphates on various response involving only a few clones of lymphocytes.
enzymes. oligoclonal bands Two or more discrete bands on electrophoresis
nucleotide Phosphate ester derivative of a nucleoside, the phos- that are produced by a few clones.
phate group (or groups) being attached to the sugar. Nucleotides oligodendrocyte Myelinating glial cell of the vertebrate central
are the monomer units of DNA and RNA. nervous system.
nucleus (1) The large and usually prominent organelle in a oligogenic Describes a trait controlled by several genes, one of
eukaryotic cell that contains the cell’s chromosomal DNA, and which usually has a major effect.
is the site of DNA replication and transcription and RNA oligohydramnios Reduction in the volume of amniotic fluid
processing. The nucleus is bounded by a double membrane that is around the fetus to below 100 ml. Usually the diagnosis is made
a continuation of the endoplasmic reticulum. (2) Anatomically by ultrasound scan and the finding of a deepest pool of less than
discrete collection of functionally related nerve cell bodies in the 2 cm is taken as diagnostic.
central nervous system. oligomer A molecule formed by connecting several (2, 3, 4, 5)
NUd See nonunion distal. smaller molecules (monomers), usually in the form of a chain or
NUp See nonunion proximal. network.
null allele An allele that produces either no protein product or a oligonucleotide A short chain of nucleotides, often made using a
nonfunctional protein product (under the conditions analysed). DNA synthesizer to create primers that can be used in PCR and
sequencing applications.
nullizygous The absence of a particular gene product as a result of
oligonucleotide hybridization probing A method of detecting
the presence of two null alleles at a locus.
particular DNA sequences in a mixture. After dissociation into
numerical aperture (NA) The sine of half the intake angle of a
single strands, the DNA is hybridized with a labelled, single-
lens, multiplied by the refractive index of the medium between
stranded oligonucleotide complementary to the target sequence.
lens and object.
oligonucleotide primer A short piece of single-stranded DNA,
nutritional ecology The study of the interrelationships between
generally 17–30 nucleotides in length, that is used in PCR and
food resources in the environment and the consumptive use of
DNA sequencing methods to start DNA replication at a
these food resources by animals.
particular sequence.
nystagmus Rhythmic oscillation of the eye during vestibular or
oligosaccharide A series of monosaccharides (from 2–20) linked
full-field visual stimulation. Each oscillation is composed of a
together by glycosidic bonds through their hydroxyl groups into
slow, compensatory and a fast, anti-compensatory phase.
linear or branched polymers.
oligotrophic Describes an aqueous environment with low levels of
nutrient supply.
O antigen A branched polysaccharide portion of lipopolysacchar- ommatidia (singular ommatidium) The individual units in an
ide that extends out from the bacterial surface. arthropod compound eye, each containing a crystalline ‘lens’, a
observational study A study of whether an exposure (a measure set of photoreceptor neurons and support cells.
at recruitment to the study, such as increased cholesterol omnivorous Having a broad diet of both plant and animal origin.
concentration) is associated with an outcome (e.g. myocardial omnivory Having a diet that includes both plants and animals.
infarction) at follow-up. Humans are traditionally categorized as omnivores.
ochre mutation Mutation that produces TAA, a nonsense codon oncogene activation The conversion of a proto-oncogene into an
that will signal the termination of translation. oncogene, caused by viral insertion, viral transactivation or
ocular dominance columns Alternating, regularly spaced, bands through gene mutation.
of left- and right-eye inputs in layer 4 of the primary visual cortex. oncogenesis The generation of a tumour, which initially involves
oedema Swelling due to abnormal collection of fluid in the the conversion of a cell to a tumour cell by, e.g., activation of
extracellular spaces of tissues. It manifests as a boggy swelling oncogenes and/or loss of tumour suppressor genes as a result of
and excessive weight gain. In the context of haemolytic disease of mutation.
the newborn, oedema occurs because of cardiac failure due to oncology The branch of medicine that deals with cancer, literally
anaemia. the science of new growths.
OH Abbreviation of the prefix ‘Olduvai Hominid’. oncoprotein The protein product of a viral or cellular oncogene.

211
Glossary

ontogenetic stage A morphologically distinct stage in develop- optic chiasm Region behind the eyes where some retinal axons
ment. cross from one side of the brain to the other.
ontogeny The growth and development of an individual from optic lobes Well developed regions of the brain concerned with
embryo to adult. processing visual data.
ontology The branch of philosophy that is concerned with basic optic tectum The visual centre in the brain of lower vertebrates,
questions about reality and existence. such as fish, amphibia and birds.
onychomycosis Fungal infection involving the nails. optical path length The effective distance travelled by a light
Onychophora A phylum/subphylum of tropical and subtropical beam in going from one point to another, allowing for the
carnivorous lobopodial arthropods inhabiting the litter layer. repertoire end view of the materials through which it passes, and
oocyst The cystic stage of a parasite. the possibility that it is scattered and does not follow a straight
oocyte An immature egg cell. line path.
oocyte maturation The resumption of meiosis in an oocyte at the optical section An image produced by some form of noninvasive
completion of oogenesis to produce the mature egg. In many light microscopical imaging and containing little or no informa-
species, the egg then undergoes a second meiotic arrest. tion from above or below the focal plane of interest.
oogenesis The development of a female egg cell in the mother, order (1) Unit of taxonomic classification that comes between
from the immature oocyte up to the mature egg. family and class. (2) A very high taxonomic unit used in
oogonium (plural oogonia) A female gametangium that contains taxonomic studies of prokaryotes.
one or more discrete gametes, the oospheres. Ordovician period The second major division of the Palaeozoic
era.
Oomycetes Also known as water moulds. Genetically, these are
ORF See open reading frame.
quite unique fungi, generally now classified separately from the
organelle A subcellular structure that performs a specialized
Fungi, that are commonly plant pathogenic and lack cross walls.
function. Examples are the nucleus, mitochondrion, endoplasmic
oosphere A large naked mass of protoplasm that functions as the
reticulum, Golgi body.
female gamete, or egg, in an oogonium.
organizer General term for the embryonic structure formed during
oospore A thick-walled sexual spore that develops from the
gastrulation which has the ability to induce a secondary neural
oosphere after plasmogamy.
tube when placed into an ectopic site. In the mouse, the node,
open complex A complex of RNA polymerase and a promoter in
or anterior condensation of the primitive streak, possesses
which the duplex DNA is unwound around the transcription start
‘organizer’ activity.
point.
organomegaly Abnormal enlargement of any or all visceral
open reading frame (ORF) A set of codons (triplets in DNA) organs.
uninterrupted by a stop codon and potentially translatable into a organometallic Involving a direct metal–carbon bond.
protein. origin of replication Site on a DNA molecule at which replication
operator Site in bacterial DNA to which a regulatory protein binds is initiated. Bacterial chromosomes and plasmids have a single
to exert its effect on gene expression. origin of replication; most eukaryotic chromosomes have
operon In bacteria and archaea, a series of adjacent genes that are multiple origins.
cotranscribed into a single mRNA, and are thus coordinately origin of transfer Sequence on a plasmid that serves as the starting
controlled. An operon comprises a regulatory gene, an operator point for the synthesis of a DNA molecule which can be
region, a promoter region and the structural genes. The protein transferred to a recipient bacterium during conjugation.
products of these genes are almost always involved in the same Ornithischia The ‘bird-hipped’ dinosaurs. These were exclusively
metabolic pathway. herbivorous and are recognized by a bird-like arrangement of
opines Low-molecular weight molecules produced by Agrobacter- their hip bones. Paradoxically, they are not closely related to
ium-transformed cells and used by the bacterium for its growth. living birds.
opioids The opiates (derivatives of opium) and synthetic narcotics Ornithuromorphs A diverse group of birds including all living
resembling opiates. birds.
opisthobranch molluscs, opistobranchs A subclass of the class oropharyngeal washing Garglings. The fluid recovered from
Gastropoda (Opistobranchia) comprising marine molluscs in washing the mouth and throat by keeping the fluid in motion
which the shell is much reduced or absent. from air expelled from the lungs.
opisthosoma The posterior body region of chelicerates. It is orthodox seeds Seeds whose viability is best maintained by dry
composed primitively of 12 segments and houses the heart, storage at temperatures close to or below 01C.
respiratory organs, reproductive system and much of the orthograde Having upright body posture.
digestive system. orthologous genes The name given to the same gene or DNA
opportunistic infection An infection caused by an organism that sequence when they occur in different species as a result of an
usually does not cause disease in individuals with a normal ancestor containing that gene giving rise to two or more different
immune system. species.
opsonin Any protein (e.g. vertebrate antibodies, complement orthologous sequence See orthologous genes.
fragments, and various factors in invertebrates) whose binding orthology (1) The situation in which similarities between two
to a foreign antigen facilitates the antigen’s recognition and species are the result of a common evolutionary origin, and in
subsequent elimination by phagocytic cells. which the gene product or structure at issue retains the same form
opsonization A process by which phagocytosis of a foreign and function. (2) The relationship between any two homologous
antigen is facilitated by its coating with opsonins (e.g. antibody, genes that are descended from the same gene in the nearest
complement fragments). common ancestor.

212
Glossary

orthorhombic Crystal symmetry group typified by three mutually outgroup In phylogenetic studies, a taxon used to establish the
perpendicular axes of unequal length. root of a phylogenetic tree. It is chosen from among ancestral
orthostatic Related to the upright position of the body. groups or species that diverged from the group being studied
oseltamavir A neuraminidase inhibitor that acts as an anti- prior to diversification within that group.
influenza agent. outgrowth The growth of a germinated spore as a vegetative
oscillation Any measurable behaviour that is periodic in time. That organism capable of synthesizing RNA, protein and DNA.
is, the state of the system repeats at regular intervals called the outside-in signalling Signalling responses that are triggered
period of the oscillation. The reciprocal of the period is the within a cell following the binding of integrins to their ligands.
frequency. overdominance The case when the heterozygote (e.g., Aa) has
osmolality A measure of the osmotic pressure generated by a higher fitness than either of the homozygous genotypes (e.g., AA
solution. It is the concentration of osmotically active molecules, and aa), thus maintaining polymorphism. That is, the reproduc-
i.e. those that cross semipermeable membranes in solution, tive success of Aa ensures that both the A and a alleles are
measured in osmoles (the standard unit of osmotic pressure) per preserved.
litre of solution. ovipary Type of reproduction in which eggs are laid and embryos
osmoregulation Regulation of a cell or organism’s water and ion develop outside the mother’s body.
balance. ovulation Process by which an ovum, or egg, is released from the
osmotic concentration The number of dissolved particles (ions ovary.
and molecules) in 1 litre of water. ovule The organ in which the female gametophyte develops in
osmotrophy, osmotrophic Heterotrophic nutritional mode in plants. In angiosperms ovules arise from part of the inner surface
which an organism acquires food material by taking it in (by of carpels and are enclosed by them. After fertilization and
active transport) through the cell membrane in a dissolved state. development of the embryo, the ovule matures into a seed.
It is characteristic of many microorganisms (e.g. several major oxidase A haem enzyme that accomplishes the four-electron
groups of internal symbiotic or parasitic protozoa). reduction of dioxygen to water.
osteoarthritis A degenerative joint disease characterized by oxidation A chemical reaction that results in the loss of electrons
erosion of articular cartilage, which can be either primary or from the molecule that undergoes oxidation.
secondary to trauma or other conditions. oxidative burst (1) Rapid production and accumulation of
osteoblast Cell that secretes bone matrix and initiates bone reactive oxygen species in infected plant tissues following
mineralization. pathogen attack. (2) The production of reactive oxygen species
osteoclast Large multinucleate cell, related to the macrophage, in macrophages and neutrophils after pathogen ingestion.
that degrades bone matrix. It is found wherever existing bone oxidative damage Damage caused to biological molecules by free
tissue is being removed and migrates over the bone surface, radicals or other reactive oxygen species generated in the normal
creating irregular scalloped cavities. course of oxygen-dependent metabolism.
osteocyte An osteoblast after it has been trapped in bone matrix oxidative pentose phosphate pathway A metabolic pathway by
and ceases to deposit bone. It is well connected to a dense network which glucose 6-phosphate is oxidized to finally yield triose
of other osteocytes and bone-lining cells through cytoplasmic phosphates and reducing power in the form of NADPH.
extensions which radiate outward from the central vascular canal. oxidative phosphorylation Formation of ATP from ADP and
osteoderms Bony dermal scutes covered by epidermal scales that inorganic phosphate (Pi) by an ATP synthase, which is coupled to
make up the carapace of armadillos and glyptodonts and are also oxidation of substrates (e.g. NADH) via the electron transport
present in mylodont sloths. system with the consumption of oxygen.
osteogenesis The production of new bone. oxidative stress Cellular stress caused by an excess of reactive
osteoid Unmineralized bone matrix, deposited by osteoblasts and oxygen species (free radicals). It occurs when there is a serious
consisting primarily of type I collagen. As the matrix mineralizes, imbalance between the generation of reactive oxygen species in
the collagen infrastructure provides tensile strength. vivo and the activity of antioxidant defences.
osteon Complex of concentric lamellae, or plates, of bone oxyanion A negatively charged oxygen atom, usually arising from
surrounding a canal containing blood vessels, nerves and loose attack on a carbonyl group. The oxygen atom receives electrons
connective tissue. from the double bond upon attack on the carbon, resulting in
osteopetrosis An inherited disease that is characterized by an excess electrons.
increased density and apposition of bone due to a reduced activity oxygen debt The amount of lactate accumulated during a bout of
of osteoclasts (macrophage-like cells that degrade bone matrix). activity, which after activity has stopped is reconverted to
osteoprogenitor cells A population of cells with the latent glucose, a process that requires an increase in the consumption
capability to produce bone-forming cells on demand. of oxygen.
ostiate heart A heart with pores to permit the entrance of blood oxygenase A haem enzyme that accomplishes addition of one
from the haemocoel. oxygen atom or two oxygen atoms to a substrate.
ostracods A group of small, mostly aquatic crustaceans that have
minute, bean-shaped, bivalved, hinged usually calcitic shells.
Phylum Arthropoda, subphylum Crustacea, class Ostracoda. P element A transposable element found in Drosophila.
They have been present from Cambrian to Recent. P granules Aggregations containing RNA and protein which are
otolith ‘Ear stone’, a structure containing calcium carbonate found in the germline and in early embryos of Caenorhabditis
particles embedded in a gelatinous mass, which sits above the elegans, where they are segregated to the future germline at each
hair cells in the maculae of the inner ear. of the early cleavages.

213
Glossary

P site The tRNA-binding site on the ribosome where the peptidyl- paracrine circuit effect Occurs when the effect of a hormone or
tRNA (P) is located before peptide-bond formation. It involves cytokine is exerted on a cell population that differs from the
both the large and small subunits. secreting cells themselves.
p53 A regulatory protein of the eukaryotic cell cycle. It acts at one paracrine growth factor A secreted polypeptide that is made by
of the checkpoints for detecting DNA damage and is an one cell and acts on a different neighbouring cell.
important tumour suppressor protein. paracrine Describes the local action of a cytokine or hormone on
PAC P1-phage derived artifical chromosome. Large-insert cloning cells close to the producer cell.
system retaining features of both the Escherichia coli F factor and parallel fibres Nonmyelinated axons of granule cells forming
the P1 phage. The vector carries an antibiotic-resistance gene, massive parallel bands running through Purkinje cell dendrites in
copy-number control elements and a positive selection system for the cerebellar cortex. Each Purkinje cell receives synapses from
cloned inserts. 100 000 parallel fibres.
pachytene The stage of meiosis I prophase during which bivalent paralogous genes Homologous genes that have arisen through
formation is completed and chiasmata appear. the process of gene duplication.
packaging signal A motif in a retroviral RNA genome that paralogy The relationship between any two homologous genes
determines their efficient encapsidation into virions. descended from the same ancestral gene by gene duplication and
packing defect Localized defects or voids in the bilayer structure divergence.
caused by the presence of membrane lipids with cis double bonds. paramutation The heritable modification of the expression state
These form kinks in the hydrocarbon chains which prevent of an allele (the paramutable allele) which depends on the
effective close-packing of the lipid hydrocarbon chains in the presence of another allele (the paramutagenic allele) that has
bilayer. acquired a repressed state.
PACS See Picture Archiving and Communication System. Paranthropus A genus introduced by Robert Broom in 1938 to
paedomorphosis One form of heterochrony, in which the juvenile accommodate a group of early hominins distinguished by
characters of the ancestral form are retained in the adult of the relatively small incisor and canine teeth, large premolars and
descendant. molars, and a wide, flat face.
PAGE See polyacrylamide gel electrophoresis. paranemic joint molecule A DNA strand-exchange intermediate
Palaeognathae The taxon of large running birds that cannot fly, where the individual complementary strands of DNA are base
often called ratites. They have a rhynchokinetic skull and a paired but not intertwined.
specific palate-reinforcing construction. The group includes the parapatric Describes populations or species with geographical
extinct moa, the ostrich, kiwi, rhea, emu and cassowary. distributions that are contiguous but not overlapping.
palaeontology The study of ancient life from fossils. paraphyletic group A group of species that includes their single
palaeosol Fossilized soil. common ancestor but not all of this ancestor’s descendants. Thus,
palatal detachment An uncoupling of some lower facial and basal members of a paraphyletic genus, for example, can also be closely
skull bones from the braincase, during evolution, through which related to members of other genera.
slenderized bones at the ventrorostral region of the skull formed a paraphyly See paraphyletic; paraphyletic group.
movable apparatus that was capable of operating the prokinetic paraplegia Paralysis of the lower extremities.
or rhynchokinetic movement of the skull. paraprotein Any unknown band of restricted migration seen on
palindrome A DNA segment in which the sequence is the same on electrophoresis, prior to identification. Once identified, the band
one strand read 50 to 30 as on the other strand. is then referred to by its specific name.
pallial Situated in the mantle cavity. paraptry See parapatric.
palpation Examination performed by touching and probing with parasporal crystal A crystalline protein inclusion formed within
the hands and fingers. the sporangium during sporulation of certain bacteria.
PAMPs See pathogen-associated molecular patterns. paratenic host A host that carries a parasite but does not support
panbiogeography A methodology for studying the origins of its growth or reproduction.
closely related species that tries to reconstruct the events leading paratope Single binding site within the V regions of an antibody or
to observed distributions of species by connecting the known T-cell receptor that binds to a particular epitope on a particular
distributions of related taxa by drawing lines on maps. antigen.
pancreatitis Inflammation of the pancreas. paraxenology Similarity arising by a combination of gene transfer
pandemic A worldwide epidemic of a disease. and gene duplication/multiplication.
panmictic Randomly interbreeding. parenchyma (1) Plant cell type with thin-walled cells and living
papilloedema Swelling of the optic nerve head visible on fundo- cellular contents at maturity. (2) The tissue occupying the space
scopic examination reflecting increased intracranial pressure. It between the epidermis and the gastrodermis in planarians. It
results from increased pressure in the nerve sheath slowing the provides support to the internal organs, and is primarily
axoplasmic transport within the optic nerve. composed of two cell types: the neoblasts and the large, highly
para-aortic Situated beside the aorta. interdigitated, fixed parenchymal cells.
parabasal body Organelle of various shape and size in flagellate parenchymella A sponge ‘larva’ composed of an envelope of
protozoans, sometimes wound about the axostyle. It is identical flagellated cells surrounding an internal cell mass.
to the Golgi apparatus and forms an identifying feature of the parental imprinting In relation to genetics, the phenomenon that
Parabasalea. for some genes the paternal copy or the maternal copy only is
parabiosis Linking pairs of animals together so that they have a expressed in the early embryo, as a result of epigenetic
common blood circulation. modification in the gamete. Also known as genomic imprinting.

214
Glossary

parts Variables expressed as parts or percentages of a whole. Parts pelagic Living in the water column, not attached to rocks or the
are dependent on each other; if one is omitted, all remaining parts seabed.
are affected. This is not so for normal variables. pellicle A set of cortical structures that strengthens the outermost
pascal The SI unit of pressure or stress (1 Pa 5 newton (N) m2), layer of the cell body in ciliates and some flagellates, enabling
commonly expressed in units of kPa and equivalent to 100 milli- them to maintain shape. A typical pellicle comprises the plasma
bars or 7.5  103 mmHg (101 kPa 5 1 atm 5 760 mmHg). membrane, two alveolar membranes and a granular cytoplasm
PAS See periodic acid Schiff technique. (epiplasm).
passerine Describes birds of the Order Passeriformes (includes pelta Crescent-shaped structure composed of microtubules and
52% of bird species), which have feet adapted for perching on associated with the axostylar capitulum in flagellates, it may be an
branches. Three-quarters are songbird families in the Suborder extension of the axostyle wrapped around the basal body.
Oscines, characterized by a versatile syrinx (vocal organ). penciclovir Nucleoside analogue inhibitor of human herpesvirus 1.
passive antibody Antibody that is present is young animals and is penetrance A measure of the ability of an allele to produce a
derived from the mother, either in utero or through the milk. The phenotype in the population.
animal’s own immune system is not stimulated. penetration The stage of viral replication at which the virus
passive transport Transport of molecules across a membrane genome enters the cell.
down a concentration or electrochemical gradient without the use penicillin-binding proteins (PBPs) Enzymes located in the
of other cellular energy. bacterial cell membrane and responsible for the terminal stages
passive-margin basin The basin at the edge of a subsiding of bacterial peptidoglycan (cell wall) synthesis. They are usually
continental margin as it moves away from the mid-ocean ridge. transpeptidases and carboxypeptidases and are the critical targets
It is usually filled with shallow marine shelf sediments and deep- for b-lactam antibiotics.
marine sediments of the continental slope and rise. pentacyclic triterpenoid A compound, containing five rings,
patch testing The placing of small amounts of a potential allergen which is made up of three C10 molecules known as terpenes.
on the skin to determine which agent might be responsible for an These compounds are distinct from, though related to, sterols.
allergic contact hypersensitivity reaction. pentose phosphate pathway A metabolic pathway in which
paternalism Interference with or overriding of another’s prefer- carbon dioxide is produced directly from glucose. A major
ences or choices in order to avoid harm to or benefit the one whose function of the pathway is the provision of NADPH.
preferences are interfered with. pentraxins A family of plasma proteins involved in innate
pathogen Any disease-producing agent, especially a virus, bacter- immunity. They are composed of five identical subunits.
ium or other micoorganism that is expected to cause disease in a PEP:PTS See phosphoenolpyruvate: carbohydrate phototransfer-
normal host. ase system.
peptide Any chain of fewer than 100 amino acid residues. Peptides
pathogen-associated molecular patterns (PAMPs) Unique
may be synthesized de novo or derived from enzymatic cleavage of
molecular patterns expressed by microorganisms which are
proteins.
recognized by the innate immune system.
peptide backbone That part of a polypeptide or protein structure
pathogenesis-related protein A diverse groups of proteins
that includes the amide bonds and a-carbons but not the
synthesized de novo in response to pathogen infection. They
individual side chains.
associate with induced defence responses and often possess
peptide nucleic acid (PNA) A synthetic DNA analogue contain-
antimicrobial activity.
ing a (2-aminoethyl) glycine backbone which forms highly stable
pathognomonic Specific and characteristic of a given disease or
duplexes with complementary DNA.
condition, as, e.g., Aschoff bodies are of rheumatic carditis.
peptidergic (1) Describes a neuron that secretes a peptide
pathway The succession of different steps in a process.
neurotransmitter. (2) Having an action resembling that of a
patristic distance Sum of branch lengths connecting two nodes in peptide hormone. The term describes a receptor that is activated
a phylogenetic tree. or modulated by a peptide.
PBDs See peroxisome biogenesis disorders. peptidoglycan The complex polymer that forms the major
PBPs See penicillin-binding proteins. structural element of bacterial cell walls. It is composed of short
PCR See polymerase chain reaction. glycan chains crosslinked by short peptides such as murein and
p-distance For DNA and amino-acid sequences, the fraction of pseudomurein.
sites at which two sequences differ in content. peptidylprolyl isomerase An enzyme that catalyses the cis/trans
pectates Pectic polysaccharides containing a high proportion of isomerization of proline residues.
1,4-linked galactosyluronic acid residues that are not methyl- peptidyltransferase Enzymatic activity that transfers an amino
esterified. acid to the carboxyl end of a growing polypeptide chain by
pectic polysaccharides Plant polysaccharides composed predo- formation of a peptide bond. Peptidyltransferase (transpeptida-
minantly of 1,4-linked galactosyluronic acid residues. tion) activity resides in large ribosomal subunit RNA, not in a
pectins Pectic polysaccharides containing a high proportion of protein.
methyl-esterified 1,4-linked galactosyluronic acid residues. peptidyltransferase centre Catalytic site on the large ribosomal
pedal nerve The large nerve running from the body of a crab into subunit where the bond between tRNA in the P-site and the
each leg. carboxyl end of the polypeptide is broken, and transferred to the
pedicle (1) The narrow stem of an organ or tumour. (2) The aminoacyl end of the aminoacylated A-site tRNA.
appendage commonly protruding from the larger of the two peptidyl-tRNA hydrolase Enzyme that catalyses the hydrolysis of
valves of a brachiopod, and generally responsible for attachment all N-blocked aminoacyl-tRNAs with the exception of fMet-
of animal to substratum. tRNAfMet.

215
Glossary

perforin A pore-forming protein present in the granules of permeability A measure of how well an ion moves across a cell
cytotoxic T cells. membrane. Membrane permeabilities for different ions are
perfusion The blood flow through tissue. usually expressed as ratios.
pericardial effusion Fluid collecting in the sac-like membrane permeance A measure of the permeability of a membrane to gases,
around the heart (pericardium) due to anaemia and cardiac liquids or solutes with units of m s1 (calculated from the flow rate
failure. across the membrane in g m2 s1 divided by the appropriate
pericarp The wall of a fruit. It develops from the ovary wall. concentration gradient in g m3).
pericentric inversion A chromosomal inversion that includes the Permian period The last major division of the Palaeozoic era.
centromere. permineralization The conversion of soft or hard tissues of an
periclinal (1) Describes a plane parallel to the outside of the organism to mineral.
organism or organ. (2) Describes a cell division in which the plane permissive interactions Conditions required for the expression of
of division is oriented parallel to the surface. a new phenotype gained through directive interactions.
pericyclic reactions A large group of organic reactions the peroxins Proteins encoded by the PEX genes and involved in the
stereochemistry of which is controlled by the conservation of biogenesis of peroxisomes. At least 20 are known and include
orbital symmetry. cytosolic and membrane receptors, putative components of the
peridinin Accessory carotenoid pigment present in many dino- membrane translocation apparatus, and many proteins of
flagellates. unknown function.
peroxisomal targeting sequence type 1 (PTS1) A C-terminal
perinatal mortality Death that occurs just before or just after
tripeptide based on the SKL amino-acid motif. This is the
birth. In humans, the term is used for stillbirths after 24 weeks or
most common targeting sequence found in peroxisomal
168 days of gestation and deaths within the first 7 days after
matrix proteins. PTS1 proteins interact with the import receptor
delivery.
Pex5p.
periodic acid Schiff technique (PAS) A staining technique in
peroxisomal targeting sequence type 2 (PTS2) An N-terminal
which material containing vicinal glycol groups on their amino or
sequence based on the XnRLX5H/QLXn amino-acid motif.
alkylamino derivatives can be oxidized by periodic acid to form
PTS2s are less common than PTS1s and unlike PTS1s are often
dialdehydes that combine with Schiff reagent to form an insoluble
cleaved after import. PTS2 proteins interact with the import
magenta compound.
receptor Pex7p.
periosteum The highly vascularized and innervated fibrous
peroxisome An organelle in eukaryotic cells that is involved the
connective tisssue layer surrounding a compact bone. It is rich
breakdown of peroxides, especially hydrogen peroxide, oxidation
in collagen, osteoblasts and fibroblasts and can be a site of new
of amino acids and lipid metabolism. It is usually characterized by
bone formation.
the presence of catalase.
periostracum Thin, organic external layer of a shell.
peroxisome biogenesis disorders (PBDs) Human genetic dis-
Peripatus See Onychophora. eases in which peroxisomes do not form properly. In many cases
peripatric speciation Formation of a new species in small the mutant genes have been cloned and shown to encode
peripheral isolated populations or beyond the periphery of the peroxins.
parent species’ range. persistent transmission The transmission of a pathogen through-
peripheral circulation The collective name for the blood vessels out the life cycle of the vector.
that supply blood to the body tissues. It is also termed the PEV See position-effect variegation.
systemic circulation. Peyer’s patches The organized lymphoid tissue of the small
peripheral membrane proteins Proteins that are bound to intestine.
membrane surfaces (usually to integral membrane proteins) but PFU See plaque-forming unit.
that do not have any of their mass buried in the membrane. PFU assay Plaque-forming unit assay. It is used to determine the
peripheral nervous system (PNS) Part of the nervous system infectivity of viruses that can infect and kill cultured cells.
encompassing peripheral nerves, sensory and sympathetic gang- PGA See 3-phosphoglycerate.
lia, and nerve roots. phage See bacteriophage.
periplasm The region between the inner and outer membrane in a phage k A temperate phage that infects the bacterium Escherichia
bacterium. It contains a large number of transport proteins and coli.
enzymes and is the region in which the peptidoglycan is located. phage vector Modified phage genome that is capable of replicat-
Extracellular bacterial proteins fold in the periplasm. ing in its host organism and into which foreign DNA (e.g. human
periplasmic enzymes Enzymes that function in the space between DNA) can be inserted for cloning.
the inner and outer membranes of Gram-negative bacteria. phagocytes Scavenger cells that take up microorganisms and cell
periplasmic region See periplasm. debris by sweeping the plasma membrane around the object and
peripodal cells Cells in the imaginal discs of insects. internalizing it in a structure called the phagocytic vacuole
perissodactyl mammals Hoofed mammals with an odd number or phagosome. In vertebrates, the ‘professional’ phagocytes
of functional toes on each foot, such as horses, tapirs, and involved in host defence are the monocytes, macrophages and
rhinoceros neutrophils, but other cells have a limited ability for phago-
perisynaptic glial cells Glial cells anatomically associated with cytosis. In invertebrate host defence, the plasmatocytes and
chemical synapses. These are astrocytes in the central nervous granulocytes are the phagocytic cells.
system, Bergmann glial cells in the retina and perisynaptic phagocytose To actively engulf large particles by the process of
Schwann cells at the neuromuscular synapse. phagocytosis.

216
Glossary

phagocytosis The engulfment of particulate material by a cell, by pheromones Secreted compounds that carry information about
surrounding the particles with the plasma membrane and an individual to other members of the same species, in which they
internalizing them in a membrane-bounded vacuole (a phago- induce a specific response.
some). pH A measure of the active acidity of a solution defined as the
phagolysosome A phagosome fused with lysosomes. negative logarithm of the H 1 ion activity (or concentration).
phagosome A membrane-bounded cytoplasmic vacuole formed PH See phenylalanine hydroxylase.
around a particle ingested by phagocytosis. pilus (plural pili) See fimbriae.
phagotrophic Describes a type of heterotrophic nutrition in which phloem The nutrient-conducting tissue of vascular plants, com-
particulate food is engulfed in a food vacuole, as in many posed of thin-walled living cells.
protozoa. Often whole organisms (usually other protists or phonophobia An intense dislike of sound which commonly
bacteria), serve as prey. accompanies the headache of migraine.
Phanerozoic The entire span of time since macroscopic fossils first phoresy A condition where a nematode or other potential parasite
appeared. rides on the body of an animal but does not harm it. It is used as a
phantom limb pain Painful sensations referred to an amputated means of distribution to new habitats.
limb. Phoronida, phoronids A phylum of tube-dwelling, worm-like
pharmacodynamics The relationship of the pharmacokinetics of animals having a horseshoe-shaped crown of tentacles (the
a drug to its biological effects, including, in the case of lophophore) around the mouth.
antimicrobial agents, the nature, speed and completeness of the 3-phosphoglycerate (PGA) Three-carbon molecule that is the
antimicrobial effect. first three-carbon stable intermediate of the C3 carbon dioxide
pharmacokinetics The behaviour of a drug in terms of its fixation pathway in plants.
absorption, distribution, binding, metabolism and elimination. phosphatase An enzyme that removes a phosphoryl group from
pharynx (1) In nematodes, a muscular pumping organ at the front another molecule.
of the digestive tract. (2) The portion of the gut between the phosphate overplus Accumulation of polyphosphate in bacteria
mouth and midgut in chelicerates. It is lined with cuticle and on transition from phosphate-poor to phosphate-rich conditions.
supplied with constrictor and dilator muscles for transporting phosphodiester bond The covalent linkage –O–P(O2)–O–
and processing food. between two nucleotides in a nucleic acid strand.
phase II trial An experimental test programme that checks the phosphodiesterase Enzyme that catalyses the hydrolysis of
efficacy of a drug on patients with actual disease symptoms. phosphodiester bonds.
phase separation Formation of a heterogeneous state of mem- phosphoenolpyruvate:carbohydrate phosphotransferase
brane lipid owing to the transition of high-melting point lipids system (PEP:PTS) A membrane transport system in prokary-
from the fluid phase to the gel phase at low temperature, resulting otes that catalyses the coupled transport and phosphorylation of
in the loss of native function of the biological membrane. a number of sugars. The transported sugar is phosphorylated
phase space A conceptual space in which each axis corresponds to during transport by a specific kinase, with phosphoenolpyruvate
one of the state variables, and in which the system is represented as the phosphate donor. This system is also involved in the
by a point at any given moment of time. chemotactic response to these sugars.
phase variation The change in antigen specificity demonstrated by phosphoenolpyruvate-dependent sugar phosphotransferase
some bacteria, caused by their on–off switching of a flagellar system See phosphoenolpyruvate:carbohydrate phosphotrans-
protein gene. ferase system.
phase–response curve The graphical representation of how a phosphoenzyme An enzyme with a covalent bond between a
phase of a rhythmic process changes when perturbed by an nucleophile on the enzyme and a phosphorylating agent.
external stimulus at different times or phases in its cycle. phospholipase C (PLC) Enzyme that hydrolyses the membrane
phasins Small, amphipathic proteins found at the boundary layer phospholipid phosphatidylinositol 4,5-bisphosphate to produce
between an insoluble inclusion body and the soluble cytoplasm of inositol 1,4,5-phosphate and diacyglycerol.
the bacterial cell. phospholipids Amphipathic lipids in which a glycerol phosphate
phenetic Describes a grouping of organisms founded on overall backbone is substituted with two fatty acids and a hydrophilic
similarity, based on all available characters, which may include base such as choline. They are the major constituents of the lipid
genotypic and phenotypic data. bilayer of biological membranes.
phenocopy An instance of a trait that has arisen environmentally phosphorite Sedimentary rock containing calcium phosphate,
but simulates a genetically determined trait. chiefly as the mineral apatite. Also called phosphatic nodule.
phenoloxidase An enzyme that oxidizes o-diphenols to phosphorolytic Describes the degradation of starch to glucose
o-quinones, eventually producing melanin. In insects it can be phosphate by starch phosphorylase.
present in both the haemolymph and the cuticle and participates phosphorylase An enzyme that uses the oxygen of phosphate as
in wound healing and sclerotization. Also called tyrosinase. the nucleophile in a substitution reaction.
phenotype The morphological, biochemical and behavioural photoautotroph, photoautotrophic Organism that utilizes the
characteristics of an individual organism or cell. Phenotype is energy derived from sunlight to build up its organic molecules
determined by both genetic and environmental factors. from inorganic materials available in their surroundings, using
phenotypic plasticity The capacity of a given genotype to produce carbon dioxide as the carbon source.
different phenotypes in response to different environmental photobiont The symbiotic alga or cyanobacterium (i.e. the
conditions. photosynthetic partner) in a lichen.
phenylalanine hydroxylase (PH) Enzyme that catalyses the photodiode A semiconductor diode that produces a photovoltage
conversion of the essential amino acid phenylalanine to tyrosine. as a result of the absorption of photons.

217
Glossary

photoheterotrophic growth Growth using organic compounds phyllody Transformation of normal floral parts (petals) into leaf-
as nutrients and light as a source of energy. like structures.
photoinhibition Reduction in the capacity for photosynthesis phyllosphere The external surface of plant leaves.
caused by high light levels, which damage the photosynthetic phyllotaxis The relative positioning of the leaves (and other above-
apparatus. ground organs) on the shoots.
photolithotroph Organism that uses sunlight as the energy source phylogenetic Relating to the evolutionary history of a species, a
and inorganic materials as the source of the elements needed for taxonomic group, or a structure.
growth, using carbon dioxide as the carbon source. phylogenetics A branch of systematics that concerns the uncover-
photolithotrophy See photolithotroph. ing of the evolutionary history of organisms and the relationships
photomorphogenetic Pertaining to the regulation of morpho- between different groups.
genesis by light, other than by the provision of organic matter by phylogenetic marker Homologous molecule whose nucleic acid
photosynthesis. or amino acid sequence information may be used to infer the
photoperiod The period of time during which a plant is exposed to phylogeny of organisms.
daylight in the context of the day–night cycle. With respect to the phylogenetic species concept The idea that a species is the
initiation of flowering, different plant species respond to different smallest diagnosable cluster of individual organisms within which
length of photoperiod. there is a parental pattern of ancestry and descent.
photophobia Intense dislike of light which commonly accompa- phylogenetic tree A branched, (usually) bifurcating representa-
nies the headache of migraine. tion of the evolutionary relationships between taxa. They can be
photophosphorylation The formation of ATP from ADP and PI based on DNA sequence comparisons or on fossil, anatomical or
driven by the energy of sunlight in photosynthetic organisms. other similar evidence.
photoreceptor neurons The light-sensitive neurons of the retina. phylogeny The evolutionary history of species and the genetic
They contain specialized structures – rhabdomeres – that absorb relationships between them.
light and transduce the energy to a signal that is sent through the phylum (plural phyla) A major taxonomic grouping of organisms
axons to the underlying optic lobes of the brain. (e.g. Arthropoda, Mollusca), based on a common body
photorespiration The biochemical reactions initiated by the organization.
oxygenase activity of Rubisco, which utilize atmospheric oxygen physical map An ordered arrangement of loci of genes and DNA
and serve to recycle some of the reduced carbon fixed by markers on a chromosome, based on the number of nucleotide
photosynthesis. pairs between loci.
photosynthate The carbohydrate products of photosynthesis. physician-assisted suicide The act of aiding a patient in suicide by
photosynthesis The physicochemical process by which some providing effective means such as drugs.
organisms (plants, algae and some bacteria) can utilize the energy physiognomy The shape of the body, as expressed, for example,
of sunlight to power the biosynthesis of organic molecules, using in indices that relate the lengths of the two limbs to that
carbon dioxide as the carbon source. of the trunk, and to each other, and in indices that reflect
photosystem A pigment–protein complex that uses light energy to the relative lengths of the component segments within each
excite specific chlorophylls that then transfer electrons to nearby limb.
cofactors, thus setting in motion photosynthetic electron trans- phytanyl Branched-chain hydrocarbons (approximately 20
port. carbons) which are highly methylated and are ether-attached
phototherapy Ultraviolet illumination of the skin of a baby to to the polar headgroups of the lipids in archaeal mem-
convert unconjugated bilirubin, released from haemolysis of red branes.
blood cells, to the conjugated water-soluble form, which will not phytoalexin A low-molecular weight antimicrobial compound
cross the blood–brain barrier. produced as a secondary metabolite by plants. It is synthesized
phototroph, phototrophic See photoautotroph. and accumulated in response to infection or other physical or
phototroph Any organism whose principal or only source of chemical stresses.
primary energy is sunlight. phytohormone An organic substance that in minute quantities
phototrophic Describes an organism that relies on photosynthetic profoundly affects plant growth. Examples include auxins,
fixation of carbon dioxide as its sole source of carbon. gibberellins and ethylene.
phycobilins Water-soluble blue or red pigments that function in phytosteroid A plant-produced molecule having the ring system
the capture of light energy. The energy is ultimately transferred to of cholesterol.
chlorophyll a. phytosterol A plant produced molecule having a cholestane
phycobiliprotein Blue/green or red water-soluble accessory pig- skeleton and alkyated at C24 in the side chain.
ments present in the chloroplasts of some algal groups. pili (singular pilus) Nonflagellar, long filamentous structures on
phycobilisome A light-harvesting structure found in cyano- the surface of bacteria. Also called fimbriae.
bacteria. It contains allophycocyanin, phycocyanin and phyco- pilin Protein subunit of bacterial pili or fimbriae.
erythrin. pinacoderm The outer epithelial layer of the body wall of a
phycoerythrin Protein involved in collection of light for photo- sponge.
synthesis in cyanobacteria. pinacocyte A cell of the sponge pinacoderm, the cell layer
phyletic gradualism A theory that suggests that most evolution separating the sponge from the external milieu.
occurs at a phyletic level, that is, within established species. ping-pong mechanism Catalytic mechanism in which one sub-
Speciation is regarded as a relatively rare event which results from strate is bound by an enzyme, transformed to product, and
a gradual divergence in morphologies. released before the second substrate is bound.

218
Glossary

pinocytosis ‘Cell drinking’. The ingestion by a cell of small plasmablast A proliferating B lymphocyte at a stage before final
droplets of extracellular fluid by endocytosis. differentiation into a plasma cell. Plasmablasts can migrate,
pituitary gland Gland situated in the base of the skull, divided into contain large amounts of intracellular immunoglobulin and
a posterior part derived from nervous tissue and an anterior part potentially also secrete it.
from glandular tissue. It secretes several hormones involved in plasmalemma The plasma membrane of plant cells. It lies
reproduction. immediately under the cell wall and invaginates and proliferates
pixel A two-dimensional picture element which represents the to surround all intrusive hyphal structures produced by mycor-
smallest digital element of an image. rhizal fungi.
PKA See protein kinase A. plasmanyl ether lipids Glycerophospholipids (primarily of the
PKC See protein kinase C. choline-linked family) in which position 1 of the glycerol
pK The pH at which an acid is half dissociated. backbone is linked via an ether linkage to a long-chain
placental abruption Bleeding into, and rupture of, the attachment alcohol.
of the placenta to the wall of the uterus. plasmenyl ether lipids Glycerophospholipids (primarily of the
placental viviparity Bearing of young nourished within the ethanolamine-linked family) in which position 1 of the glycerol
maternal uterus by transfer of nutrients through a placenta. backbone is linked via a vinyl ether linkage to a long-chain
Placentalia, placentals The name of the clade stemming from the alcohol.
most recent common ancestor of all modern ‘placental’ mammals plasmids Extrachromosomal DNA present in bacteria and yeast
(all mammals excluding the marsupials and monotremes). Also and capable of replication independently of the chromosome.
known as the Eutheria. Most are in the form of a supercoiled circular double-stranded
placentamegaly Enlargement of the placenta. DNA, but some linear plasmids are known. Plasmids are used in
placode A cake-like thickening of the ectoderm that gives rise to recombinant DNA procedures as carriers for foreign genetic
sensory neurons and, in the case of the inner ear and the lateral material.
line, to hair cells. plasmodesmata (singular plasmodesma) Narrow channels
placoid Tooth-like scale structure typical for sharks. through plant cell walls which provide cytoplasmic continuity
planktic Describes aquatic organisms that passively float. between adjacent cells, enabling them to communicate with each
plankton Tiny animals and plants that live suspended in the water other and to exchange metabolites.
column, with little or no contact with the bottom. They drift in the plasmodium (plural plasmodia) The multinucleated single-cell
ocean and provide food for many marine animals. feeding stage of plasmodial slime moulds and some protostelids.
planktonic Describes aquatic organisms that are floating passively In the plasmodial slime moulds, this cell can be very large and
or only weakly swimming. features the fastest rate of cytoplasmic streaming known in
planktotrophic Describes actively swimming larvae that feed nature.
within the plankton. plasmogamy The fusion of cytoplasm of two protoplasts.
planktotrophic larva Larva feeding in the plankton. plasmolysis Shrinkage of the plant cell protoplast due to loss of
plant growth regulators Natural and synthetic compounds that water, forming a space between the plasma membrane and the cell
modify plant growth and development. wall.
plant segmentation Independent responses of different parts of a plasticity (1) The property of the potential for change or
plant to external conditions. This can be due to localized cambial modification. In life sciences, the term is used, for example, to
activity and to vascular conduits favouring one organ over describe the capacity of a phenotype to vary according to
another. environmental experience, and to describe the ability of the brain
planula Pear-shaped to rod-like solid embryo of cnidarians, which to make and reinforce new connections between neurons in
develops into a tubular polyp. response to experience. (2) Term used to describe the ability of a
planum temporale An area of the brain adjacent to the primary molecule to undergo structural changes without extreme input of
auditory cortex of the temporal lobe. The left planum temporale energy. (3) The tendency of a material to change shape in response
is relatively larger in individuals who have absolute pitch. to an applied load.
plaque A small clear area that forms around an initial focus of plastids Organelles in plant and algal cells that have a characteristic
phage infection on a lawn of susceptible bacteria (or of a virus on double membrane or envelope. Examples are chloroplasts and
susceptible cells), due to lysis of infected cells. With an chromoplasts.
appropriate concentration of phage (or virus) in the initial plastochron, plastochron ratio The time interval between the
inoculum, each plaque will be due to a single initial phage (or initiation of successive leaf primordia.
virus) particle. plastocyanin A soluble copper-containing protein found in the
plaque-forming unit (PFU) An infectious virus or phage particle thylakoid lumen. Plastocyanin carries electrons from the
that can undergo replication in an infected cell and then reinfect cytochrome bf complex to photosystem I.
surrounding cells to form an area of cell death termed a plaque. plastoquinone A small organic molecule involved in electron
plasma cell dyscrasia Proliferation of a single, usually malignant, and proton transfer in photosynthesis. Plastoquinone is
clone of plasma cells. Characterized by production of large found in the hydrophobic core of the thylakoid membranes
quantities of identical immunoglobulins. and in the photosystem II reaction centre and cytochrome bf
plasma membrane The lipid–protein bilayer that surrounds the complex.
cytoplasm. Also called the cell membrane, cytoplasmic mem- plate tectonics The movement of segments of the Earth’s crust as a
brane, plasmalemma (in plants) or inner membrane (in Gram- result of its production at mid-ocean ridges and burial in ocean
negative bacteria). trenches.

219
Glossary

platelet A small membrane-bound enucleate cell fragment (de- poikilotherm A ‘cold-blooded’ organism whose body temperature
rived from megakaryocytes) present in large numbers in blood is generally very close to environmental temperature. Elevation of
plasma (typically 2–2.5  108 per ml). When exposed to extra- body temperature may be produced by behavioural means such
vascular components of injured tissue it releases factors that as basking, but not via biochemical reactions of heat production.
induce cell division and blood clotting. poikilothermy Maintaining a variable body temperature equal to
platelet-activating factor A molecule that induces blood platelets that of the surroundings.
to release factors that induce cell division and blood clotting. It is pointed end The slow-growing end of an actin microfilament.
a 1-alkyl phospholipid with an acetyl group in the 2-position. point mutation A change in a single base pair in the coding or
plating efficiency The proportion of phage particles that, when noncoding region of a gene.
plated on a lawn of susceptible bacteria, will yield a plaque. Poisson correction A transformation applied to raw p-distance
platyrrhine A general term of primate classification used for the measurements to correct for multiple substitutions at a site.
New World monkeys. polar filament A thin multilayered tube made of protein and coiled
PLC See phospholipase C. around the internal periphery of the microsporidian spore. It is
plectonemic joint molecule A DNA strand-exchange intermedi- thought to evert when the spore germinates, thus pulling the
ate where the complementary strands of DNA are both base sporoplasm through and injecting it into the host cell.
paired and intertwined. polar pili Protein filaments located at the extremities of a
pleiotropic Describes a gene or mutation that affects more than bacterium.
one phenotypic characteristic. polar tube See polar filament.
pleiotropy The multiple effects of a single gene on different organs, polarity (1) The direction of change of a character, from one state
or organ systems. to another. (2) Having one end morphologically or biochemically
pleocytosis The presence of an abnormal (increased) number of distinct from the other. (3) Situation in which electrical charge is
cells in the spinal fluid. not distributed uniformly in an atom or molecule.
pleomorphic Exhibiting more than one distinct form. Having polarizability Response of a molecule’s electron density to an
morphologically distinct stages in the life cycle. applied electric field or the reponse of a material to an applied
pleopods Abdominal arthropod appendages often used for swim- field.
ming. polaroplast A highly membranous structure attached near the
base of the polar filament and located behind the anchoring disc
plerology Similarity that arises as a result of gene rearrangement,
at the anterior of the microsporidian spore.
either within or between genes.
pollen The mass of male gametophytes, or pollen grains, of a seed
plesiomorphic Describes an ancestral character, in an evolution-
plant, which are produced within the anther. Each pollen grain
ary sense.
contains three cells: two sperm and a vegetative cell that grows a
plesiomorphy The original (primitive) state of a character.
pollen tube in order to deliver sperm to embryo sac.
pleurae (singular pleura) The lateral parts of the trilobite’s
poly(A) signal DNA sequence transcribed as part of an mRNA
thoracic segments, on either side of the upraised axis.
that determines the formation of the 30 end of the mRNA and the
plexus A network of vessels or nerves. In the latter case the nerves
addition of the polyadenylate tail.
may split and exchange nerve fibres.
poly(A) tail A homopolymer of adenine ribonucleotides that is
ploidy The number of chromosome sets present in a cell or added to the 30 end of eukaryotic mRNAs after transcription.
organism. Haploids have one chromosome set; diploids have two; polyacetate A polyketide metabolite synthesized only from an
and polyploids have more than two. acetate starter and n-malonyl extenders.
plurilocular Describes multicellular gametangia and sporangia, polyacrylamide gel electrophoresis (PAGE) A technique for the
each cell of which produces a single motile cell, which functions as separation of molecules from complex mixtures (proteins and
a gamete or asexual spore, respectively. nucleic acid fragments) by their differential migration in an
pluripotent Describes a cell that can give rise to a number of electric field in a polyacrylamide gel. It is typically used to
different cell types. separate DNA fragments between 10 and 1500 bp long.
pluripotent stem cell A stem cell that gives rise to many different polyadenylation The post-transcriptional addition of a string of
types of terminally differentiated cells. nontemplated adenine nucleotides to the 30 end of an mRNA
plus end The end of a microtubule that is fastest growing. It is the molecule.
natural end for the assembly of tubulin subunits, the end towards polyandry The situation where females mate with several males,
which the b subunits of tubulin dimers are directed, and the end usually within the same reproductive period.
towards which most kinesin motors translocate. Poly-b-hydroxyalkanoate granules Intracellular deposits of
plus end, minus end The fast and slow-growing ends, respectively, alkanoate polymers formed in some bacteria.
of a microtubule (occasionally also used to refer to the barbed and Poly-b-hydroxybutyrate granules A type of polyhydroxyalk-
pointed ends of actin filaments). anoate granule.
pluton A body of magma that has cooled at depth in a magma polycistronic Describes a bacterial messenger RNA containing the
chamber. coding sequences for more than one protein.
PNA See peptide nucleic acid. polycistronic transcript An mRNA molecule resulting from the
PNS See peripheral nervous system. continuous transcription of several adjacent genes.
PNS See postnuclear supernatant. polyclonal Composed of several different clones of cells.
poikilohydric Describes organisms that require high humidity but polyclonal antibodies The mixture of antibodies usually evoked
tolerate desiccation and rehydrate when favourable conditions by a single antigen, which contains antibodies that recognize
resume. different epitopes on the antigen.

220
Glossary

polycythaemia An increase in the proportion of cellular compo- polyphyletic group A group of species that does not include their
nents of the blood with an inevitable decrease in the plasma single, common ancestor.
component, which may have adverse effects. polyphyletic origin Describes a trait that has been independently
polydentate binding An association between different molecular established in a number of different lineages.
species in which noncovalent interactions (binding) take place at Polyplacophora The chitons, a class of the phylum Mollusca.
several sites in each molecule. polyploid (1) Possessing three or more sets of chromosomes. (2)
polyembryony The formation of more than one embryo in the ovule. Having more than one copy of the virus genome per virus particle.
polygenic Caused by or involving many genes. polyploidy The condition in which there are three or more sets of
polygonum-type embryo sac The most common type of embryo chromosomes in a cell.
sac in higher plants (named after the plant Polygonum divaricatum). polyprotein A long polypeptide which is composed of the
It is an eight-nucleate embryo sac derived from a single spore. sequences of several different proteins, which are subsequently
polygyny The situation where males mate with several females, released by protease cleavage.
usually within the same reproductive period. polysaccharide A polymer of more than 20 monosaccharide units
polyhedral bodies Inclusions possessing many faces. joined into a linear or branched chain through glycosidic bonds.
polyhydramnios An abnormal condition of pregnancy when there polysaccharide-protein A compound of plant origin in which one
is excess production of amniotic fluid in the uterus. or more polysaccharide structures are covalently attached to a
polyketide A secondary metabolite formed by a common bio- polypeptide chain.
synthetic pathway involving decarboxylative condensation of polysialic acids Natural polymers of N-acetylneuraminic acid
malonyl units. (sialic acid).
polymer A large molecule formed by connecting many chemically polysome A messenger RNA molecule with two or more translat-
similar small molecules (monomers) together, usually in the form ing ribosomes attached.
of a chain or network. Proteins (polypeptides) are polymers of polysomnography A formal study of sleep, which is used to detect
amino acids. Nucleic acids (DNA and RNA) are polymers of obstructive sleep apnoea.
deoxyribonucleotides and ribonucleotides respectively. Polysac-
polyteny The condition when a chromosome is composed of many
charides are polymers of sugars.
identical DNA duplexes, which is caused by many rounds of
polymerase In the most general sense, an enzyme that catalyses the
DNA replication not followed by separation.
reactions that connect molecular subunits to one another to form
polytypic species A species constituted of two or more locally
polymers. Most commonly refers to any of the enzymes that
adapted populations (subspecies), with little gene flow among
catalyse the assembly of ribonucleotides or deoxyribonucleotides
them.
into RNA (transcription) or DNA (replication). DNA poly-
polytypism The existence of two or more genotypically/pheno-
merases catalyse the production of DNA molecules. RNA
typically distinct populations of the same species that have been
polymerases catalyse the production of RNA molecules.
recognized as geographic races or subspecies.
polymerase chain reaction (PCR) An in vitro method for
poly(U) A polynucleotide composed of uridine nucleotides only.
specifically amplifying a given sequence in a DNA molecule.
polymorphic Existing in many different forms. polyuria–polydipsia Combination of excessive thirst (polydipsia)
polymorphism (1) The existence of a character, or a gene, in two or and excessive urine output (polyuria).
more forms in a population, where the least common form is population A group of organisms of the same species occupying
present in more than 1% of individuals. (2) A region of DNA the same area and isolated to some degree from other popula-
sequence that differs from allele to allele. tions.
polymorphs Polymorphonuclear leucocytes or neutrophils, population bottleneck A period during which a population is of
phagocytic, antimicrobial white blood cells with irregularly greatly reduced size and genetic drift plays a larger role than usual
shaped nuclei. in changing allele frequencies.
polymorphonuclear leucocyte See neutrophil. population doublings The estimated number of times the cell
polymyositis polyneuropathy Any systemic disorder of the number in a culture has doubled.
peripheral nerves of toxic, metabolic or unknown aetiology. population viability analysis The use of mathematical models to
polypeptide A linear polymer of amino acids covalently linked to determine the likely viability of a population in the future.
one another through bonds called ‘peptide bonds’. Geneticists population-specific allele See private allele
tend to distinguish between a polypeptide, by which they mean pore Conductive pathway through e.g. a membrane, which is
the linear chain of amino acids, and a protein, in which the chain is generally always open.
folded into a three-dimensional conformation. porphyria A disorder of haem metabolism that can be either
polyphenism The situation when an organism is able to produce genetically transmitted or induced by certain chemicals.
two or more distinctive phenotypes in response to environmental portal vein A blood vessel that joins a set of capillaries in one organ
cues. It contrasts with polymorphism, which is based on genetic with another so that substances can pass directly between the two
differences. organs.
polyphosphate granules (bodies) Intracellular deposits of con- position effect variegation (PEV) The silencing of a gene in some,
densed phosphates. but not all, of the cells in which it is normally expressed, which
polyphyletic Having distinct evolutionary histories. The term e.g. gives rise to a variegated appearance of the structure. The
describes a group of organisms that are classified together, often silencing is due to a chromosomal rearrangement that brings the
because of convergent evolution, but are derived from a number gene under the influence of heterochromatin.
of different ancestral species, so that they are not all descended positive chemotactic stimulation An increasing gradient of
from a common ancestor that was a member of that group. chemoattractant or a decreasing gradient of chemorepellent.

221
Glossary

positive eugenics Programmes, techniques or social policies that power Rate of energy use or production or of doing work. It is
encourage and favour reproduction among individuals believed measured in watts (W) (= joules (J) per second (s)).
to carry desirable genes. ppGpp Guanosine 30 ,50 -bis(diphosphate). An ‘alarmone’ synthe-
positive feedback The type of control in a metabolic or signalling sized by bacteria to overcome amino acid starvation and involved
pathway in which a late stage in the pathway has a reinforcing in growth rate regulation.
effect on early stages. ppm Parts per million, a measure of concentration.
post ribosome The post-translocational state of a ribosome, with pRb See Rb
deacylated tRNA in the E site and the peptidyl-tRNA in the P preacquisition fasting period Fasting of aphids for a defined
site. period prior to allowing acquisition of virus. Such a period is
postcranial That part of the skeleton of vertebrates that is behind, mostly needed for nonpersistent viruses.
or beneath, the cranium. It includes the vertebral column, both pre-B cell A stage in the development of B cells that occurs between
limb girdles and all four limbs. the pro-B cell and the immature B cell. Pre-B cells express a
posterior Towards the tail end of an organism. The anterior- complete m heavy chain but have not yet rearranged the light-
posterior axis of a vertebrate organism, for example, runs from chain genes.
the tip of the nose, snout or beak to the tip of the tail. Prebilateria A group of primitive animal phyla that do not display
posterior fossa The posterior and inferior portion of the cranium bilateral symmetry and are believed to have evolved before the
that contains the brainstem and cerebellum. bilaterians.
posterior vacuole A large vacuole at the posterior of the precursor cells Cells that are committed to a particular cell lineage
microsporidial spore. It is thought to be the part of the extrusion but are still in a process of developing the characteristics of that
apparatus responsible for pushing the cytoplasm out of the spore cell type.
during germination. precursor frequency The frequency of a lymphocyte subset with a
postganglionic sympathetic Describes neurons of the peripheral given antigen specificity.
autonomic nervous system normally projecting from the para- pre-eclampsia Pregnancy-specific syndrome associated with new
vertebral or prevertebral ganglia. high blood pressure and proteinuria, appearing in the second half
postmitochondrial supernatant The fraction of a cell homo- of pregnancy and resolving after delivery.
genate that remains after removal of mitochondria and other pre-mRNA The unspliced precursor to messenger RNA.
large subcellular particles by centrifugation. prenatal diagnosis The detection of fetal abnormality during
postnuclear supernatant (PNS) Cell homogenate containing pregnancy.
membranes and cytosol after removal of nuclei by slow-speed pre-pro-B cells The earliest precursors of B cells identified so far.
centrifugation. They have DH and JH gene segments of the immunoglobulin
postpartum oestrus The oestrus that occurs immediately after genes rearranged but not the V gene segments.
giving birth. pre-ribosome The pretranslocational state of a ribosome, with
postsynaptic Refers to the cell that is on the receiving side of a peptidyl-tRNA in the A site and deacylated tRNA in the P site.
synapse. presoma The anterior portion of the two-segmented preparasitic
postsynaptic membrane The plasma membrane of the cell on the larva of nematomorphs, which contains the reversible proboscis.
receiving side of a synapse. Also called the preseptum.
postsynaptic potential (PSP) The change in membrane potential presomatic cell A cell that undergoes chromatin diminution in e.g.
in a postsynaptic (receiving) neuron or muscle cell caused by the Parascaris, and thus will become a somatic cell.
action of transmitter released from the presynaptic neuron on prespore See forespore.
postsynaptic receptors. presynaptic Refers to the cell that is on the transmitting side of a
post-temporal canal An opening on each side of the posterior part synapse.
of the skull in certain reptiles, retained in monotremes. presynaptic facilitation Enhanced transmission from a synaptic
post-tetanic potentiation (PTP) See potentiation. terminal of a neuron (the presynaptic cell) onto a target
post-transcriptional gene silencing A reduction in gene expres- postsynaptic cell that results from the action of a third neuron
sion associated with a sequence-specific degradation of RNA in (the facilitator cell) synapsing onto the axon terminal of the
the cytoplasm. presynaptic cell.
post-transfer editing Hydrolysis of misacylated tRNA with the pretransfer editing In tRNA synthetase proofreading of amino
liberation of free noncognate amino acid and tRNA. acids, the destruction of noncognate aminoacyl adenylate,
post-transfusion purpura Thrombocytopenia caused by destruc- usually by hydrolysis to free noncognate amino acid and AMP.
tion of transfused and autologous platelets, occurring about 1 prevacuole A membrane-bounded compartment in eukaryotic
week after blood transfusion and associated with the presence of cells intermediate between the Golgi and the vacuoles.
anti-platelet alloantibodies. prevalence The percentage of a host population that is infected by
post-translational Occurring after translation is complete. symbionts.
post-translational modification Any modification that occurs to primary cell wall A polysaccharide-enriched matrix that encapsu-
a polypeptide chain after its synthesis is complete. lates growing plant cells.
potentiation Slowest phase of enhanced transmission in short- primary culture A culture of unpassaged cells isolated directly
term synaptic plasticity, lasting minutes. from tissue, organs or body fluids.
potocytosis Mechanism for the cellular uptake of small molecules primary image plane An image of the object is formed at the
or solutes independent of endocytosis but mediated by caveolae. primary image plane owing to interference between those beams
pouch A fold of abdominal skin around the mammary glands of a diffracted by the specimen and the undiffracted (zero order)
monotreme or marsupial. beam.

222
Glossary

primary immune response The immune response to an initial processivity The property of DNA and RNA polymerases that
exposure to antigen. enables them, once bound, to move along the template,
primary immunodeficiency disease Immunological disorder successively adding nucleotides to the growing DNA or RNA
resulting from an inborn defect of the immune system. strand.
primary root Root that has its origin in the plant embryo. processivity error An error in translation in which peptidyl-tRNA
primary structure The sequence of amino acids along the is prematurely released. The corresponding nascent peptide is
polypeptide chain of a protein. shorter than the mature protein and is thus usually nonfunc-
primary transcript The order or sequence of nucleotides, from the tional.
50 to 30 end, within a strand of RNA exactly as it is transcribed procoracoid A bone that connects with the shoulder, the scapula
from the DNA, before processing. and the interclavicle, forming a bony bridge in the pectoral
primer (1) A short sequence of RNA synthesized on the DNA region.
template that provides a startpoint for the synthesis of a DNA procumbent Protruding anteriorly.
strand in DNA replication. After DNA synthesis is complete, the prodrome Symptoms of yawning, euphoria, unbridled energy or
RNA is removed and replaced with DNA by repair enzymes. depression, or craving or distaste for various foods, that occur in
(2) A short oligonucleotide of specific sequence that is used to the 24 hours before a migraine headache.
define the startpoint for DNA synthesis in the polymerase producer An organism, such as a photosynthetic, photolitho-
chain reaction. See also mutagenic oligonucleotide primer. trophic or chemotrophic organism, that can utilize solar energy or
primitive In the context of taxonomy, a species with relatively few energy derived from inorganic chemicals to manufacture its own
specialized features, thus indicating that it is closer to the food (carbohydrates) from carbon dioxide. Producer organisms
morphology of the earliest members of an evolutionary lineage. are the basis of all life on Earth and support the remainder of the
primitive ectoderm See epiblast. food chain.
primitive haematopoiesis Formation of blood cells that occurs proecdysis See moult cycle.
before birth or hatching, usually marked by the expression of proenzyme A precursor form of an enzyme that has not yet been
different globin genes and cell morphology. activated to its final functional form. Activation is usually
primitive life A chemical system capable of self-reproduction and achieved by cleaving an inhibitor domain from the protein.
evolution.
progametangium (plural progametangia) A swollen hypha
primitive streak A narrow elongated region of mesoderm inter- that will differentiate into the gametangium and suspensor.
nalization in early bird and mammalian embryos, indicating the
progenitor A precursor cell that has not yet undergone maturation
axis along which gut and notochord will form, and the antero-
of differentiation and maintains the ability to divide.
posterior axis of the embryo.
progenote A putative ancestral organism from which all extant
primordial germ cell A cell which is the precursor of germ cells
organisms are thought to have derived.
only.
programmed cell death A regulated mechanism of cell death by
primordial leaf A leaf in the earliest stages of its development, as it
apoptosis that eliminates cells in a predictable fashion, without
starts to form on the shoot meristem.
damage to the organism, suggestive of an underlying develop-
primordium (plural primordia) A structure in its most rudimen-
mental or genetic programme. It is common during development,
tary form, at a very early stage in its development.
e.g. in the formation of the digits of hands and feet, and in the
PRINS See primed in situ labelling.
resorption of the tadpole tail during metamorphosis.
prion Infectious proteinaceous particle that is a conformational
progress zone Mesenchyme at the distal tip of the limb. The
isoform of a normal cellular protein. Prions are thought to be
progress zone mesenchyme is kept in an undifferentiated and
responsible for some degenerative diseases of the central nervous
proliferative state by signals from apical ectodermal ridge. Call
system, such as Creutzfeldt–Jacob disease in humans, and bovine
fate specification is believed to occur in the progress zone.
spongiform encephalitis (BSE) in cattle.
prismatic Showing crystalline structure. prohaemocyte A stem cell from which other haemocyte types
originate in invertebrates. It is characterized by a nucleus that
prismatic cartilage Outer layer of cartilage in sharks, arranged in
almost fills the cell and the absence of intracytoplasmic granules.
crystalline prisms.
private allele A genetic variant, evolved by mutation in a specific prokaryotes The members of the Bacteria and the Archaea. They
population, which did not get a chance to spread to other are small single-celled micoorganisms (about 1–2 mm in diameter)
populations of the species. whose DNA is not enclosed in a nucleus and in which the
cytoplasm is not compartmented into membrane-bounded
probe (1) A short oligonucleotide (DNA or RNA), usually
organelles.
labelled, that is used to detect the complementary sequence in a
DNA by hybridizing with it. (2) A fluorescently or radioactively prokaryotic Pertaining to prokaryotes.
labelled antibody or other protein that is used to label a cellular prokaryotic cells See prokaryotes.
structure or particular molecule in a sample. (3) A fluorescent Prolacertiformes An important group of Triassic reptiles closely
molecule whose emission is used to analyse a sample. related to the archosaurs, the group that gave rise to the
procambial initial A precursor cell in the procambium that can dinosaurs. They were generally lizard-like in body plan and
differentiate into various types of vascular cells including commonly had a narrow skull and long snout.
tracheary element, xylem parenchyma cell, cambial cell, sieve proliferation Increase in the number of cells by means of mitotic
element and phloem parenchyma cell. cell division.
procambium Cells destined to become the primary vascular tissue promastigote The stage of Leishmania that occurs in the insect
in plants. vector.

223
Glossary

promiscuity In the molecular context, the tendency of a molecule protein b sheet A secondary structure in proteins that is formed
to bind to many alternative ligands rather than to have a strict when two or more almost fully extended polypeptide chains are
specificity for only one other molecule. brought together side by side so that hydrogen bonds form
promoter A region of DNA that is typically located immediately between the peptide amide NH and carbonyl oxygen groups of
before a gene and that contains signals for initiating transcription adjacent chains.
of the gene. It contains the binding site for RNA polymerase and protein binding motif An amino-acid sequence that is present in
its ancillary factors. several proteins that can interact with an RNA molecule.
pronephros The first embryonic kidney in vertebrates, maintained protein disulfide isomerase An abundant enzyme in the endo-
only in some fishes. plasmic reticulum that catalyses disulfide bond formation and
proneural Describes genes whose activity pushes cells towards a isomerization during protein folding.
neural fate. protein domain A compact structurally discrete portion of a
proofreading In the context of translation, the ability of protein, usually with a particular function.
aminoacyl-tRNA synthetases to prevent or correct mistakes in protein electrophoresis A technique used to separate different
the aminoacylation of tRNA. forms of a protein on the basis of their different rates of
pro-oxidant Stimulating or catalysing oxidative reactions poten- movement in response to an applied electric field through a liquid
tially damaging to cells. or a gel.
propagule pool colonization The founding of a new protein family A group of proteins that are suspected to be
population with colonists derived from a single source evolutionarily related on the basis of their amino acid sequence or
population. folding pattern.
prophage The bacteriophage genome in a state when it is not protein folding The process by which a linear polypeptide chain
producing new phage particles. The genome may be integrated spontaneously folds into its functional three-dimensional native
into the DNA of the host bacterial cell or be replicating as a conformation.
plasmid. protein kinase An enzyme that transfers phosphate groups from
ATP to proteins, usually phosphorylating the protein at serine,
prophenoloxidase An enzyme precursor stored in granulocytes
tyrosine or threonine residues, depending on the type of kinase.
and which produces a functional phenoloxidase when activated
protein kinase C (PKC) An intracellular enzyme that is activated
by enzymatic cleavage.
by a variety of signal transduction events and functions to
prophylaxis Prevention of disease, for example, by antibiotic
phosphorylate specific proteins.
administration before surgery.
protein module Any protein domain or protein sequence that is
prosimian A general term of primate classification that includes
used repeatedly as a building block in diverse, mostly extra-
the lemurs, lorises and the tarsier, even though tarsiers are more
cellular, proteins.
closely related, evolutionarily, to anthropoids.
protein phosphorylation The addition of a phosphate group to
prosoma The anterior body region of chelicerates. It is composed protein, usually catalysed by a protein kinase.
of six appendage-bearing segments and is typically covered by a
protein synthesis The biosynthesis of a protein on a ribosome by
dorsal carapace bearing the eyes.
the linking together of amino acids in a sequence directed by a
prosome The first body section of a phoronid, represented by the messenger RNA.
epistome in the adult and by the preoral lobe in the actinotroch protein targeting The allocation of a protein to a specific region
larva. or organelle within the cell, or for secretion. The targeting is
prospective study See observational study. mediated by specific sequences within the protein.
prostaglandins A class of derivatives of the unsaturated fatty acid protein tertiary structure The overall spatial arrangement of
arachidonic acid, formed via the cyclooxygenase pathway. They amino-acid residues within a single folded polypeptide chain.
are involved in many biological processes, including the contrac- proteinase inhibitors Small peptides, rather resistant to degrada-
tion of smooth muscle, the development and control of tion, with inhibitory activity towards proteinases (proteases).
inflammatory responses, the regulation of body temperature, protein-tyrosine kinase An enzyme that phosphorylates a protein
and the sensing of pain. at tyrosine residues.
protandry Ambisexuality where the individual functions first as a protein-tyrosine phosphatase An enzyme that cleaves phos-
male, then as a female. phate groups from tyrosine residues in a protein.
protease An enzyme that can cleave a protein at a peptide bond. proteinuria The presence of excessive amounts of protein in the
Some proteases cut nonspecifically, catalysing the breakdown of urine.
proteins to peptides and amino acids; others selectively cut at a Proteobacteria The largest and physiologically most diverse
certain combination of amino acids. group of bacteria, grouped into five subclasses by probable
proteasome Large catalytic multisubunit protease located in the sharing of ancestry with phototrophic purple bacteria.
cytoplasm which degrades cytosolic proteins. proteoglycan A protein to which long glycosaminoglycan (carbo-
protected areas Areas managed primarily for the protection of hydrate) chains are attached. The carbohydrate frequently
species, biological communities and other aspects of biological accounts for much of the net mass of the molecule.
diversity. proteolytic cleavage Trimming of proteins or peptides into
protein A large organic molecule made up of one or more smaller fragments by specific enzymes, i.e. proteases.
polypeptide chains folded into the appropriate three-dimensional proteolytic processing The process by which one or more smaller
configurations and sometimes modified by the addition of protein molecules are generated from a larger precursor by site-
carbohydrate units, phosphate groups, or other chemical specific cleavage catalysed by a protease.
adducts. proteome All the proteins expressed by a cell.

224
Glossary

Protista, protists Unicellular, coenocytic or simple multicellular pseudogamy A form of apomixis in which meiosis does not occur,
eukaryotic organisms that are not classified as fungi, animals or but development of the embryo requires fertilization of the
plants. central cell by a sperm from pollen.
protogny Ambisexuality where the individual functions first as a pseudoknot See RNA pseudoknot.
female, then as a male. pseudomurein An archaeal analogue of peptidoglycan consisting
proton electrochemical potential (DmH1 ) A measure of the of alternating N-acetylglucosamine and N-acetyltalosaminuronic
potential (typically in mV) across a biological membrane due to a acid linked by b1,3 bonds. It occurs in Methanobacterium,
proton (H1 ) gradient. It is composed of a membrane electrical Methanobrevibacter and Methanosphaera.
potential (Dy) and an ion concentration gradient (DpH). pseudomurein Cell-wall peptidoglycan of some methanogenic
proton gradient An unequal distribution of H1 ions (protons) on Archaea that lack muramic acid.
both sides of a membrane. pseudomycelium A group of budding yeast cells in a tree-like
protonephridium (plural protonephridia) Excretory organ in formation of branched chains.
larva of Kinorhyncha, Phoronida and Entoprocta. It is closed at pseudopersistent transmission Type of transmission of viruses
its inner end and opens to the exterior by the nephridiopore. found in some nematode vectors. it resembles stylet-borne
proto-oncogene A gene that encodes a protein required for transmission, but with long retention times, giving a false
normal cell survival and/or proliferation, but which can give rise impression of persistent transmission.
to an oncogene after mutation or activation. When the mutant pseudopodia (singular pseudopodium) Mobile finger-shaped,
gene was originally found in a transforming retrovirus, the name lobose or net-like projections extended from the surfaces of
of the cellular form of the gene may be prefixed with a ‘c-’, and the certain cells, e.g. some protozoa, and typically associated with
viral form prefixed with ‘v-’ (e.g. c-src and v-src). shape change and movement resulting from cytoplasmic flow
protoperithecium The female sexual organ of certain ascomy- within the projection.
cetes. pseudorevertant Descendant of a mutant in which the negative
protoplast A cell including the plasma membrane but excluding effect of a mutation has been neutralized by a compensatory
the cell wall. mutation at a site in the genome different from that of the initial
Protostomia, protostomes Animals that are characterized by mutation.
certain patterns of development, e.g. formation of the mouth at pseudotypes Virus particles produced during replication of
the location of the primary invagination (the blastopore). They heterologous viruses in the same host cell in which the genome
include Arthropoda (insects, crustaceans, etc.), Annelida (earth- of one virus is contained within the coat or envelope of another
worms, polychaetes, etc.) and Mollusca (clams, snails, etc.). virus.
protostomy Describes a pattern of development in which the site pseudouridine (Y, W) 5-Ribosyluracil, a C-glycoxyl isomer of the
of the primary invagination in the early embryo becomes the classical N-glycosyl nucleoside, uridine (1-ribosyluracil). It
mouth. contains the only C–C ‘base–sugar bond’ in RNA and is the
prototroph A bacterium (or other microorganism) that has no most abundant modified nucleotide in RNA.
specific nutritional requirements for growth. PSP See postsynaptic potential.
prototrophy A phenotypic state in which a particular nutrient psychrophiles Microorganisms whose optimum temperature for
does not need to be supplied for growth. A prototrophic strain growth is below 201C. Some psychrophilic algae can grow near
resembles the wild-type strain with respect to its ability to grow 01C.
without a particular nutrient. pteridophytes Plants (e.g. ferns) in which sexual reproduction
protozoa A large, rather arbitrary, group of single-celled non- does not involve seed production, and in which there are
photosynthetic eukaryotic microorganisms lacking cell walls, and independent alternating sexual (gametophyte) and asexual
thus resembling animal cells rather than plant or fungal cells. (sporophyte) generations.
protraction Forward and/or upward motion of an anatomical pterodactyloids Advanced, short-tailed, Jurassic and Cretaceous
element relative to an overall muscle–bone apparatus. pterosaurs 0.3–12 m in wingspan.
provirus Double-stranded DNA copy of a retroviral genome PTH See phenylthiohydantoin.
integrated into host chromosomal DNA. PTP Post-tetanic potentiation. See potentiation.
pruritus (1) A condition that has as the dominant symptom a desire PTS1, PTS2 See peroxisomal targeting sequence type 1; peroxiso-
to scratch some part of the body to relieve irritation. (2) A mal targeting sequence type 2.
sensation that elicits the desire to scratch. P-type ATPases A superfamily of transmembrane proteins with
pseudarthrosis An ‘artificial joint’ that forms at the site of a ATPase activity that use the energy released by hydrolysis of ATP
fracture as a consequence of persistent nonunion of the fracture to transport ions across membranes. The enzymatic mechanism
ends. of P-type ATPases involves a phosphorylated intermediate, hence
pseudoautosomal region A region common to the X and Y the name P-type.
chromosomes in which pairing and recombination may take puberty The time when an animal becomes capable of breeding,
place. normally coincidental with the start of gametogenesis.
pseudocoelom In nematodes, the fluid-filled space separating the Pulmonata, pulmonates, pulmonate molluscs Subclass of the
ectodermal–mesodermal layer from the endodermal layer. class Gastropoda comprising snails and slugs, in which the mantle
pseudocolour A colour representation of a grey scale, defined by cavity is used as a lung.
the user. pumps Transmembrane proteins that (usually) use the energy of
pseudocospeciation A match between host and parasite phylo- ATP hydrolysis to move ions from one side of the membrane to
genies that is due to cospeciation. the other, typically against a concentration gradient.

225
Glossary

punctuated equilibrium A theory of evolution that, in contrast to radiocarbon dating A method of dating based on the decay rate of
phyletic gradualism, relates most evolutionary change to rapid carbon-14 (14C) in organic materials such as wood.
branching events, with periods of relative stasis between radiograph An image obtained using X-rays and a film (or so-
speciations. called film-screen combination).
purines, purine bases One of the two common types of bases in radioisotope A radioactive atom possessing a different number of
nucleotides. They are composed of a heterocyclic pyrimidine ring neutrons from its parent atom. In biology, radioisotopes of
combined with an imidazole ring. The purines in nucleic acids are common biological atoms such as phosphorus and sulphur are
adenine and guanine. used to label biological molecules for detection purposes or for
Purkinje cell A type of large neuron that extends characteristic use as tracers.
planar dendrites, forming a layer in the cerebellar cortex and Radiolaria, radiolarians Subclass (phylum Circadian, class Acti-
providing its sole output. nopoda) of tiny marine planktonic protozoans that secrete
purple membrane Regions of the cell membrane of halobacteria spheroidal to bell-shaped silica tests. They are important
that contain a two-dimensional crystalline array of the light- sediment formers in the deep sea. Present from the Cambrian to
driven proton pump bacteriorhodopsin, which has a purple the present-day.
colour. Radiology Information System (RIS) A computer system (in the
purpura Small bruises on the skin caused by the coalescence of past, usually mainframe types; more recently, powerful PCs) that
small pinpoint-sized skin haemorrhages called petechiae. handles workflow components of a radiology practice including
scheduling, billing and reporting.
purulent infections Infections caused by extracellular pathogens
RAG1, RAG2 Recombinase-activating genes that encode the sub-
such as streptococci or staphylococci, which cause pus-forming
units of a heterodimeric protein required for recombination of V,
lesions (abscesses) that are composed mainly of polymorpho-
D and J gene segments during antigen-receptor gene rearrange-
nuclear leucocytes and bacteria.
ment in lymphocytes.
pygidium The tail of a trilobite, often rounded or triangular in
Ran A small, Ras-related GTP-binding protein that is central to the
form, and made of fused segments like those of the thorax.
nuclear transport cycle. The yeast homologue is called Gsp1p.
pyloric caeca Blind, finger-like projections of the proximal random amplified polymorphic DNA (RAPD) DNA synthesized
intestine just beyond the pyloric sphincter of the stomach. via a polymerase chain reaction using arbitrary primers, which
pyrenoid Specialized region in the chloroplast involved in starch results in DNA fragments of varying lengths that give a specific
formation. banding pattern when separated electrophoretically.
pyridone ring Compounds based on the 5- carbon, 1-nitrogen ring random genetic drift The chance fluctuations of allele frequencies
system. Pyridones carrying iodine were used as early X-ray within a population from generation to generation as a result
contrast agents. of chance events such as sampling of gametes. The smaller
pyridoxal phosphate Chemical name for vitamin B6, found as a the population, the greater the random fluctuations in gene
cofactor in enzymes. frequency will be.
pyrimidine tract binding protein (PTB) A protein that binds to random walk A trajectory in space characterized by a sequence of
pyrimidine-rich sequences in RNA. steps in which each new step is taken in a random direction.
pyrimidines One of the two types of bases in nucleotides. They are randomized controlled trial A trial in which participants have
heterocyclic six-membered rings. The pyrimidines in nucleic acids been randomly assigned to intervention with a treatment.
are uracil, cytosine and thymine. range In palaeontology, the observed distribution of fossil finds
pyrophosphate (O)3P–O–P(O)3. describing the span between the oldest and youngest finds. If
sampling were perfect, a taxon’s range would be its duration.
range extensions Hypothesized extensions of observed ranges
implicit to particular estimates of phylogeny.
R gene A host resistance gene that specifically recognizes a gene
RAPD See random amplified polymorphic DNA.
product (Avr) of the pathogen. Note this narrow definition of an
raphe A longitudinal slit in one or both surfaces of a diatom cell
R gene, as compared to any other host gene that confers resistance
wall, necessary for movement to occur, and found only in pennate
to a pathogen and can be called a ‘resistance gene’.
diatoms.
R plasmid A plasmid containing antibiotic-resistance gene(s), rare-allele advantage See frequency-dependent selection.
which confers resistance to those antibiotics on the bacterium rate equation Differential equation describing the change of
carrying it. concentration of a reaction component over time.
R0 Index used in epidemiology, indicating the number of secondary ratite Describes a bird having a keelless sternum, the converse of
cases that could originate from a single index case of virus carinate. A member of the Ratitae.
infection. RB See Rb.
RAD52 epistasis group A group of genes, first identified in the Rb The retinoblastoma protein (Rb or RB depending on species), a
yeast Saccharomyces cerevisiae, which act in concert to achieve regulatory protein of the eukaryotic cell cycle. It is an important
homologous recombination in eukaryotic cells. tumour suppressor protein, whose absence leads to the eye
radial distribution function (rdf) Probability that one molecule tumour retinoblastoma.
or atom will be at a certain distance from another molecule or RD gene A gene situated in the class III region of the human major
atom. histocompatibility complex between the BF and C4A comple-
radiation hybrid A hybrid cell formed by fusing an irradiated ment genes.
donor with an unirradiated host cell. The hybrid retains some rdf See radial distribution function.
fragments of the donor’s chromosomes. rDNA Genes that encode the ribosomal RNA (rRNA).

226
Glossary

reaction centre, reaction centre complex A chlorophyll– receptor (1) A protein molecule that recognizes and binds another
protein complex that uses light energy to create a stable charge (the ligand) with high affinity and specificity, the binding event
separation from the excited chlorophyll or bacteriochlorophyll. usually triggering a biological response in the cell carrying the
A single electron is transferred energetically uphill from the donor receptor, although some receptors may simply act as ‘passive’
molecule to an acceptor molecule, both of which are located in the carriers. (2) A cell-surface protein that is used by a virus for
reaction centre. attachment.
reaction coordinate The single vibrational mode of a reaction receptor channel An ion channel in which a receptor site for a
assembly that has no restoring force, and hence an imaginary neurotransmitter forms an integral part of the channel itself, and
frequency. binding of the neurotransmitter is responsible for channel
reaction norm The range of possible phenotypes that a single opening, for example the nicotinic acetylcholine receptor
genotype can express along an environmental gradient. channel.
reactivation Causing something to function again when it has receptor integral anion channel An anion-selective aqueous
ceased functioning. For example, the procedure by which isolated pore formed within a receptor protein.
subcellular organelles are ‘brought back to life’, usually by the receptor-mediated endocytosis Internalization of a protein
addition of ATP. after it has bound to its receptor at the cell surface.
reactive amyloidosis Extracellular accumulation of amyloid recessive allele Allele that has to be present in two copies at a locus
protein fibrils in various organs and tissues of the body secondary for its phenotype to be expressed. In heterozygotes with a
to a chronic inflammatory disease. dominant allele, the recessive phenotype is obscured by that
reactive gliosis Strong proliferation of reactive, swollen of the dominant allele. Recessive alleles usually encode an
astrocytes accompanied by an accumulation of microglial cells inactive version of the gene product or an absence of the gene
at affected brain areas. While not specific for prion diseases, product.
gliosis is extremely prominent in Creutzfeldt–Jakob disease and recessive mutation A mutation that results in a phenotypic effect
scrapie. only when in the homozygous state. In the heterozygous state the
reactive oxygen species Collective term used by biologists normal allele prevails over the mutant allele.
to designate the oxygen radicals (e.g. O2, OH) and certain recessiveness When the phenotype of the Aa heterozygote is
nonradical derivatives of O2, such as hydrogen peroxide identical to the phenotype of the AA homozygote, the a allele is
(H2O2). said to be recessive to the A allele.
reactogenic Capable of causing an unwanted side reaction, as in
reciprocal synapse A specialized type of synapse in which the two
‘reactogenic vaccine’, e.g. causing fever, rash or excessive local
cells on either side can both transmit and receive.
inflammation.
reciprocal translocation A translocation in which part of one
reading frame In a protein-coding mRNA, the translation
chromosome is exchanged with part of a different, nonhomolo-
machinery has to start at a particular point in order to read
gous, chromosome.
off the correct sets of three nucleotides (codons) as it moves along
the RNA. There are three possible reading frames in reciprocity The situation in which an action at one point, which
any sequence, only one of which usually encodes a functional produces an effect at a second point, can be assumed to provide an
protein. accurate model of the performance at the first object of an action
realism The philosophical thesis that classes are real, contrasted at the second.
with nominalism. recirculation Continuous migration of mature lymphocytes from
realized heritability A measure of a trait’s heritability based on the blood to the secondary lymphoid organs and from there via
the trait’s response to selection over multiple generations. lymphatic vessels back to the blood.
reassociation The formation of a duplex by two nucleic acid recoding Reprogrammed genetic decoding. Three classes have
strands (DNA, RNA or both) having complementary base been identified: the meaning of specific codons can be redefined;
sequences. reading frames can be switched by ribosomal frameshifting;
reassortment The process of random redistribution of genomic blocks of nucleotides within a coding sequence can be bypassed
RNA segments into new virions following simultaneous infection by ribosomes.
of a cell by different strains. The resulting viruses are known as recoding site A sequence in an mRNA that contains special
reassortants. elements, often including a stop signal, and where an alternative
RecA protein A 38 kDa bacterial protein that plays a key role in genetic event occurs other than those characteristic of normal
homologous recombination by catalysing homologous pairing protein synthesis.
and strand exchange between the recombining DNA molecules. recognition (1) The rejection and selection of taxa that
recalcitrant seeds Seeds whose viability is lost on drying and also occurs during the establishment of a symbiosis, resulting in
on freezing. the observed specificity of a symbiosis. (2) In cell and
recalcitrant Describes a compound that resists breakdown to its molecular biology, the highly specific interactions that occur
mineral constituents. The term should not be confused with between different proteins and between proteins and DNA
xenobiotic, although they are often used with the same meaning. sequences.
For example, lignin is recalcitrant, but not xenobiotic as it is of recognition sequence DNA sequences recognized by restriction
organic origin. enzymes.
recapitulation The biogenetic law according to which the evolu- recognition species concept The idea of a species in which the
tionary history of a species (phylogeny) is repeated in a speeded- members share a specific mate-recognition system, ensuring
up form in the embryological development of the individual effective syngamy within a population occupying a preferred
organism (ontogeny). habitat.

227
Glossary

recombinant (1) Pertaining to DNA into which a foreign gene has refractory period A period following an action potential where it
been introduced by genetic engineering. (2) Pertaining to a is more difficult (relative refractory period) or impossible
protein produced by a microorganism whose DNA has been (absolute refractory period) to elicit another action potential.
engineered in order to produce it. refugia (singular refugium) Isolated areas that have served as a
recombinant DNA A single DNA molecule formed by joining refuge for a species during climatic or other changes.
together DNA molecules from different sources (e.g. genes from regenerative xylem Tracheary elements that are formed from
different parts of the same genome, or DNA from different parenchyma cells rather than from the meristematic procambium
organisms). or cambium. They may connect growing lateral buds, lateral
recombinant virus A virus that has been genetically manipulated roots, and new strands around wounds.
to incorporate a foreign gene into its genetic material. regional specification The case when a group of cells is
recombinase A protein that mediates a DNA rearrangement determined as a whole by cell–cell interactions that depend on
reaction (recombination). their position within the embryo.
recombination (1) The exchange of similar or dissimilar sequences regiospecificity The exclusiveness of modification of a specific site
between two DNA molecules. (2) Any process that gives rise to in a reactant, thereby forming a specific product when two or
cells or individuals in which parental alleles have been inherited in more sites of modification on the reactant are available.
new combinations. regulatory cascade A sequence of regulatory proteins A, B, C y
recombination mediators Accessory factors that function to where protein A catalyses the activation of protein B, which then
enhance the activity of a recombinase. catalyses the activation of protein C, and so on.
recombinational repair A DNA repair process involving an regulatory RNA An RNA whose primary function is the regula-
interaction between a damaged DNA molecule and a separate tion of a biological activity.
DNA duplex with homology to the damaged region. It proceeds regulon Groups of genes and operons that are coordinately
by reactions very similar to those of genetic recombination. controlled.
reconstitution The bringing together of separate cell components, RE See reproductive effort.
each individually inactive, but which together can carry out a reification Examination of the morphometric implications of the
given reaction. components of a latent vector by assigning biological or physical
reconstruction Process by which data acquired from multiple meaning to these.
acquisitions are manipulated to produce an image. rejection antigen An immunogenic protein in a cell that can
rectification The direction in which a cell membrane most easily induce rejection of that cell by the host’s immune system.
passes current. For example, a membrane that passes outward RelA (p)ppGpp synthetase I (PSI), an enzyme involved in the
current more easily than inward current is said to show outward ‘stringent response’ linking amino acid availability and RNA
rectification. accumulation in bacteria.
recurrent circuit A series of synaptic connections between neurons relaxed DNA Closed circular DNA without any supercoiling.
in which the last neuron connects some of its output back to the release The stage of viral replication at which virus particles escape
first. A recurrent circuit can support positive feedback. the infected cell.
recurrent flagellum A posteriorly directed flagellum. remodelling In relation to bone, a coupled sequence of resorption
recycling compartment Tubular compartment localized in the and formation events that replace previously existing bone. Also
perinuclear region or subapical region (in polarized epithelial considered as the last stage of bone healing, following the
cells), important for recycling receptors and other plasma inflammatory and reparative stages.
membrane components. Renette cell A ventrally located gland in nematodes that collects
redox The process of reduction or oxidation. liquids from the body cavity and passes them into the excretory
redox buffer A mixture of a thiol and its corresponding disulfide. duct from which they are expelled through the excretory pore.
redox reactions Reactions in which the reacting species change repair (1) DNA repair. The cellular processes resulting in the
oxidation states. removal of damaged DNA and its replacement with normal
reducing terminus The single sugar group at one end of a undamaged DNA in the genome of an organism. (2) Wound
polysaccharide chain that is not glycosidically bonded to any repair. The replacement of lost tissue by connective tissue
other group. All the other sugar groups in a polysaccharide chain elements and parenchymal cells in varying proportions. When
are nonreducing. replaced completely by granulation tissue, which later matures to
reduction A chemical reaction that results in the gain of electrons fibrous tissue, the result is referred to as a scar.
by the molecules that undergo reduction. repair synthesis of DNA DNA synthesis that fills in gaps
redundancy In respect to the genetic code, the fact that there are generated by the excision of damaged stretches of DNA.
several different codons for each amino acid. repairosome A putative multiprotein complex comprising many,
Reed–Sternberg cell The neoplastic element of Hodgkin disease. perhaps all, of the proteins involved in nucleotide excision repair
It is a large cell, either binucleate or with a bilobate nucleus, and of DNA.
possessing prominent nucleoli. repeat induced point mutations (RIP) A premeiotic process in
reflection A narrow beam of strongly diffracted X-rays produced Neurospora in which both copies of duplicated DNA are heavily
when a crystal is in a particular orientation in the X-ray beam. mutated, leading to inactivation of genes.
refractometric optics Optics based on exploiting the refractive replication See DNA replication.
index of the medium through which light is passing. The replication band Lightly staining, migrating crossband of a
ultracentrifuge employs a Rayleigh interferometer in which the macronucleus, found in all hypotrichous ciliates and in some
difference in refractive index between a reference solution and the oligotrichs. The area concerned is involved in DNA replication
sample is measured. and histone synthesis.

228
Glossary

replication fork trap Two groups of terminators with opposite resolving power In microscopy, the ability to distinguish struc-
orientations arranged to trap replication forks in the terminus tures that are close to each other (e.g. the light microscope has a
region of the chromosome. resolving power of 0.4 mm, whereas the transmission electron
replication fork Junctions in partially replicated DNA where the microscope has a resolving power of 0.2–0.5 nm).
newly synthesized daughter strands meet the unreplicated resonance a condition occurring when the driving frequency
parental DNA. The replication fork is the site at which DNA closely matches the natural frequency of the system it is driving.
synthesis occurs. Maximum energy transfer and oscillation amplitude occurs.
replication origin The point on a genome at which DNA resource defence polygyny Behaviour in which males defend a
replication initiates. necessary or limiting resource and mate with all females that come
replication protein A (RPA) A heterotrimeric single-stranded to that resource.
DNA-binding protein that plays important accessory roles in resource tracking The case where a parasite is associated with a set
DNA metabolism by preventing formation of or disrupting of hosts that share a given resource.
secondary structures within single-stranded DNA. respiration (1) Cellular oxidation of chemical fuels, leading to the
replication slippage Mispairing between DNA strands that can generation of metabolic energy and reducing power. The term
occur during the replication of tandem repeats, leading to repeat usually refers to aerobic respiration, in which the final electron
amplification or deletion. acceptor is molecular oxygen and which generates large amounts
replicative intermediate A partially double-stranded structure of ATP. (2) A type of metabolism using external electron
formed during the replication and transcription of RNA acceptors (e.g. O2).
viruses. respirators Filtered breathing devices that can protect culturists
replicative senescence An ‘ageing’ phenomenon in which normal from particles and microorganisms.
cells proliferating in culture lose their potential for replication respiratory burst Augmented oxygen consumption and produc-
and acquire biomarkers of cellular senescence after a certain tion of reactive oxygen radicals and other microbicidal products
number of serial passages in culture. which is stimulated in phagocytes (e.g. macrophages and
neutrophils) by the phagocytosis of microorganisms or by other
replisome A large multiprotein complex, located at the replication
external stimuli.
fork, that catalyses DNA replication.
respiratory chain A series of redox proteins within a membrane
reporter gene The coding sequence for a protein whose activity
(such as the inner mitochondrial membrane) that couples
can be easily assayed. It can be fused to the sequence of another
substrate oxidation to the generation of a proton electrochemical
gene of interest to report on its expression, or it can be
gradient across the membrane.
placed under the control of enhancer and promoter sequences
respiratory circulation The collective name for the blood vessels
of interest to report on their activity and tissue-specificity, for
that supply blood specifically to the respiratory organ. It is also
example.
termed the branchial or gill circulation in fish and the pulmonary
reporter molecule A molecule used in an indirect assay that
circulation in vertebrates with lungs.
detects an event that is otherwise unobservable by direct
respiratory control The tight coupling of the rate of mitochon-
examination.
drial respiration to the ratio of ATP/ (ADP + Pi) in the cell.
repressor A gene regulatory protein that shuts off the expression of response regulator In a two-component regulatory system, a
a particular gene or group of genes by inhibiting the initiation of protein, that is phosphorylated by a histidine kinase on an
transcription. In bacteria, classical repressor proteins such as the aspartate residue as part of the signal transduction process. The
lac repressor act at sequences known as operators, adjacent to the response regulators have 120 conserved residues in the N
promoter. terminus.
reproductive effort (RE) The fraction of resources allocated to restoration Returning a species to a habitat where it used to occur,
reproduction. or changing an ecosystem back to its original condition.
reproductive isolation Evolutionary independence of popula- restriction enzymes Endonucleases from bacteria that recognize
tions due to intrinsic features that prevent their successful specific DNA sequences and cut the DNA at a particular site at or
interbreeding, and thus prevent gene exchange. near the recognition sequence.
reproductive value A way of evaluating the relative demographic restriction fragment length polymorphism (RFLP) DNA is cut
importance of the different categories (age, size etc.) in a by specific restriction enzymes resulting in DNA-pieces of
population. The reproductive value of an individual in a individually varying lengths, thus giving a specific banding
particular category is the number of newborns needed to affect pattern when separated electrophoretically.
population growth as much as that individual. restriction fragment length polymorphism (RFLP) Genetic
repulsion Mechanism by which an axon or a cell is driven away. variation in a population that manifests itself in the form of
Chemorepulsion involves the interaction of the cell with soluble variability among individuals of the lengths of restriction
repellent molecules, contact repulsion involves cell–cell interac- fragments produced by cleavage of a given region of the genome
tions. by a restriction enzyme.
reservoir host Nonhuman natural host for a virus, providing a reticular formation A network of neurons and fibres occupying
continual source of the infectious agent. the core of the brainstem.
resolution (1) The ability to distinguish between events close reticulate evolution The fusing of previously separated branches
together in space, in time or in appearance. (2) The ability of an on an evolutionary tree.
imaging system to distinguish between two points in a specimen in reticulocyte A red blood cell newly made and released from the
terms of distance or time, which is often a function of the display marrow. After supravital staining, reticulocytes exhibit a
device in a digital imaging system. cytoplasmic reticular pattern detectable by light microscopy.

229
Glossary

reticuloendothelial cells, reticuloendothelial system Non- rhamnogalacturonan I A pectic polysaccharide with a backbone
migratory phagocytic cells in the marrow and lymph nodes and of alternating rhamnosyl and galacturonosyl residues. Between
the sinusoids of the liver and spleen, and macrophages in other 20 and 80% of the rhamnosyl residues may be substituted at
tissues. position 2 with oligosaccharide side chains.
retinal error Difference between fovea position and target position rhamnogalacturonan II A structurally complex pectic polysac-
in space. charide present in the walls of all higher plants. It contains several
retinoblastoma protein See Rb. unusual glycosyl residues.
retinotectal system Chick visual system with topographically rhamphorhynchoids Primitive, long-tailed Triassic and Jurassic
ordered projections from retinal ganglion cells to their immediate pterosaurs 0.5–2.5 m in wingspan.
brain target, the tectum. rhamphotheca Keratin cover of the upper and lower mandibles,
retrograde transport The movement of materials from the replacing teeth.
terminals of a neuron to the cell body. rhapidosomes Rod-shaped tubular inclusions.
retroperitoneum The area behind the rear of the abdominal rhizomorph A linear aggregation of hyphae, often robust and
cavity. root-like in appearance and sometimes showing internal differ-
retrotransposons See transposable elements. entiation, through which nutrients are transported.
retroviral packaging cell Cell line engineered to contain all the rhizosheath A sheath of soil particles bound together
proteins needed for retroviral replication. with polysaccharides that surrounds the roots of some desert
retroviruses A family of enveloped viruses with RNA genomes plants. They provide the roots with protection against desicca-
that replicate via a DNA intermediate through the use of reverse tion.
transcriptase, and in which the double-stranded DNA inter- rhombomere Segmental unit of the developing hindbrain. The
mediate may become integrated into the host chromosome as an hindbrain is composed of 7–8 rhombomeres.
inactive provirus. In this form the provirus can be passed on from rhizome A creeping underground stem from which aerial shoots
generation to generation. are produced. In ferns, the leaves and roots arise from the
reversal of damage A type of DNA-repair process in which rhizome.
a DNA lesion is repaired without excision and resynthesis of
rhynchokinetic Applied to the skulls of birds that have flexible
DNA. regions near the tip of the upper beak.
reverse genetics The term coined to describe the analysis of
rhynchosaurs A common group of large, herbivorous Triassic
gene function starting from the gene, into which mutations can
reptiles closely related to the archosaurs. They were characterized
be introduced and their phenotypic effect studied, in contrast
by broad, powerful skulls that were well adapted to grinding up
to classical genetic analysis which starts with a mutant
tough plants.
phenotype.
ribavirin A broad-spectrum antiviral drug.
reverse transcriptase RNA-dependent DNA polymerase, which
ribonuclease (RNAase) An enzyme that cleaves RNA into
catalyses the synthesis of DNA on an RNA template.
fragments or single nucleotides.
reverse transcriptase PCR (RT–PCR) Polymerase chain reaction
using as template DNA produced by reverse transcription. ribonuclease H (RNAase H) A nuclease that digests the RNA
reverse transcription The synthesis of DNA using RNA as the strand of an RNA–DNA duplex. It is found in association with
template. reverse transcriptase and is essential in reverse transcription
replication.
reversed phase high-performance liquid chromatography
(RP-HPLC) Chromatographic technique for separating peptides ribonucleic acid See RNA.
or proteins, in which the sample is run under pressure through a ribonucleoprotein motif An RNA-binding domain in a
steel column packed with 3–5 mm silica particles that carry protein that consists of approximately 80–90 amino acids
extended hydrophobic C2 to C18 chains. The particles have pore containing two highly conserved sequence elements, RNP1 and
diameters of 100 to 300 Å. RNP2.
reversed transport Transport in a direction opposite to that ribonucleoprotein (RNP) A complex consisting of one or more
occurring normally. In the case of plasma membrane amine RNA components and one or more protein components.
transporters, reversed transport moves the substrate from the cell Examples include the ribosome, telomerase, RNase P and the
interior towards the exterior. spliceosome.
reversion Restoration of a phenotypic trait to its original state, or ribonucleoprotein shedding Elimination of ribonucleoprotein
restoration of a mutated gene to its original sequence (genetic during meiotic divisions, as described in the oocytes of some
reversion). A phenotypic trait may revert as a result of a true species (e.g. Lepidoptera, Trichoptera and others).
genetic reversion or through additional, secondary alterations in ribosomal A site See A site.
the genome. ribosomal frameshifting The programmed alteration of a reading
reward conditioning See appetitive conditioning. frame by a ribosome during translation.
Reynolds number A dimensionless number that represents the ribosomal P site See P site.
ratio of inertial to viscous forces. ribosomal proteins The stable protein components of ribosomes.
RF II DNA Full length M13 bacteriophage DNA. In bacteria they are named S1–S21 for the small (30S) subunit,
RFLP See restriction fragment length polymorphism. and L1– L36 for the large subunit (50S).
RGG motif An RNA-binding motif 20–30 amino acids long that ribosomal RNA maturation The cleavage and/or chemical
is characterized by arginine–glycine–glycine (RGG) tripeptide modification of the initial rRNA transcript. The mature
repeats interspersed with other, often aromatic amino acids. rRNA is functional in ribosome assembly and protein
rhabdomyolysis Necrosis of muscle cells. synthesis.

230
Glossary

ribosomal RNA (rRNA) The non-coding RNA that is a stable RNA polymerase The enzyme that carries out transcription. It
structural component of the ribosomes. Each ribosomal RNA is a catalyses the formation of a complementary RNA strand on a
single-stranded molecule, generally denoted by its sedimentation DNA template.
coefficient, e.g. 16S rRNA, S5 rRNA. In eukaryotic ribosomes RNA polymerase holoenzyme The complete form of bacterial
the small subunit contains a 16S rRNA, and the large subunit a RNA polymerase, containing s factor, which is required to
23S and a 5S rRNA. initiate transcription efficiently. After initiation, the s factor is
ribosomal subunits The two ribonucleoprotein complexes (the lost and the rest of the polymerase, the core enzyme, is responsible
small subunit and the large subunit) that form a functional for elongation of the RNA strand.
ribosome. A large and a small subunit come together on RNA polymerase II The enzyme responsible for synthesis of
messenger RNA at the initiation of protein synthesis. The large messenger RNA in eukaryotes.
and the small subunits have sedimentation coefficients of 50S and RNA primary structure The sequence of ribonucleotides (G, C, A
30S in prokaryotes and 60S and 40S in eukaryotes, respectively. or U) in the RNA chain.
ribosome Large nucleoprotein particle, composed of multiple RNA processing The alterations that are made to primary RNA
ribosomal proteins complexed to rRNAs. Ribosomes are present transcripts in order to produce a translatable or functional RNA.
in large numbers in the cytoplasm, and are the sites at which These include RNA splicing, polyadenylation, capping, and
messenger RNA is translated to protein. methylation.
ribosome-binding site Site on bacterial messenger RNA, also RNA pseudoknot A structure in an RNA formed by a stem–loop
called the Shine–Dalgarno sequence, that is required for efficient with a tail that base pairs and forms a helix with part of the loop.
ribosome binding and translation. RNA secondary structure The three-dimensional conformation
of a folded RNA chain resulting from base-pairing between
ribozyme A polyribonucleotide with enzymatic activity, also
different parts of the chain. The base-pairing generally follows
known as an RNA enzyme. Natural examples include self-
Watson–Crick rules, but G–U pairs are also allowed.
splicing introns and the peptide-bond forming activity of the
ribosome. RNA splicing The removal of introns from a primary RNA transcript
and the rejoining of the exons to form a mature RNA molecule.
ribulose 1,5-bisphosphate carboxylase/oxygenase See Rubisco.
RNA structure Three-dimensional structure formed by intramole-
ribulose 1,5-bisphosphate (RuBP) The five-carbon compound cular interactions between the bases in an RNA molecule.
carboxylated by Rubisco in the C3 cycle. RNA tertiary structure Structure formed by superfolding and
richness The number of distinct units within a (usually taxonomic) long-range interactions in an RNA molecule.
group (e.g. species richness, generic richness) or within a habitat. RNAase See ribonuclease.
Ringer solution A ‘physiological’ salt solution made to be isotonic RNAase H See ribonuclease H.
with serum. It usually contains Na1 , Cl, K1 , Mg21 , Ca21 and RNAase P A ribonuclease involved in RNA processing that
bicarbonate, as well as lactate or glucose. matures the 50 end of tRNA transcripts.
RIP See repeat induced point mutations. RNAi See RNA interference.
RIS See Radiology Information System. RNP See ribonucleoprotein.
ritonavir Protease inhibitor of human immunodeficiency virus 1 RNP motif See ribonucleoprotein motif.
(HIV1). Robertsonian change Translocation between two nonhomolo-
RNA Ribonucleic acid. Nucleic acid, usually single-stranded except gous acrocentric chromosomes to form a bi-armed chromosome
in some virus genomes, which is composed of ribonucleotides. (Robertsonian fusion), or the mutation of a bi-armed chromo-
The bases in RNA are adenine, cytosine, guanine and uracil. See some into two acrocentrics (Robertsonian fission).
also messenger RNA, ribosomal RNA, transfer RNA. Robertsonian translocation, Robertsonian fusion Chromoso-
RNA editing A change in the information content of a messenger mal rearrangement involving two acrocentric chromosomes
RNA by the deletion, insertion or chemical modification of which fuse near the centromeric region with loss of the short
specific bases during or after transcription. These changes are arms. The result is a chromosome with two arms and a
sometimes directed by antisense RNAs. centromere in the middle.
robusticity A technical term to describe limb bones that are
RNA enzyme See ribozyme.
relatively short but which have broad shafts. Long bones are said
RNA hairpin A strand of RNA that is folded back on itself and held
to be ‘robust’ when the width of the shaft is greater than that for
together by complementary base pairing. The turning point (the
an equivalent modern human bone of the same length.
loop) contains a few unpaired nucleotides.
rod elements Arrays of aggregated chlorophyll that form the core
RNA helicase An enzyme that unwinds double-stranded RNA in of the chlorosome.
an ATP-dependent reaction. rod outer segment (ROS) A specialized compartment in the rod
RNA interference (RNAi) The phenomenon whereby a gene’s cells of the retina that consists of densely packed membrane.
function can be selectively inhibited by double-stranded RNA rolling The rotating movement of blood cells along the vessel wall
corresponding to that gene. The mechanism of this effect is still that is due to weak interactions between adhesion molecules on
unclear. the cell and the wall.
RNA maturation The complete set of enzymatic and physico- rolling-circle replication A type of replication of circular DNA or
chemical events that convert a primary RNA transcript into a double-stranded RNA in which one strand of the double-
mature functional RNA. stranded circle is nicked, and the 30 end thus generated is used
RNA modification The chemical changes that occur on individual to prime DNA synthesis, displacing a 50 tail. The single
nucleotides in RNA, e.g. change of functional groups (usually replication fork thus formed moves indefinitely around the
methylation) and isomerization. circular template.

231
Glossary

root exudates Material released by roots including sugars, amino saltation (1) The evolution of new species by sudden, discontin-
acids, organic acids, proteins, cell debris and carbon dioxide. uous steps. (2) A radical and discontinuous reorganization of
root pressure An osmotic pressure generated in the roots, pushing morphology in a single generation (e.g. through mutation),
water upwards. Thought to have important roles in young plants resulting in a population successfully occupying a new and
and in dissolving air bubbles in the xylem at the beginning of the substantially different ecological niche.
growing season. saltatory conduction The propagation of action potentials from
rosette A structure in the plant cell plasma membrane that is node to node in myelinated axons.
thought to be an enzyme complex responsible for cellulose sampling intensity The probability of sampling a taxon over some
synthesis. specified stratigraphic or temporal interval.
ROS See rod outer segment. sandfly The insect vector of leishmaniasis.
rostral Towards the snout or beak end of an animal, i.e. the anterior Santana Formation Early Cretaceous rock sequence in Brazil that
end. has produced many exceptionally well preserved pterosaur
rotavirus Double-stranded RNA virus of the genus Rotavirus, remains.
family Reoviridae, that is the major cause of infant diarrhoea. SAP See serum amyloid protein.
saprotroph An organism, especially a bacterium or fungus, that
RPA See replication protein A.
lives by absorbing nutrients from dead organic matter.
RP-HPLC See reversed phase high-performance liquid chromato-
saprotrophic Living on dead organic matter.
graphy.
saprotrophy A form of chemo-organotrophy common in bacteria
rRNA See ribosomal RNA. and fungi in which organic compounds are absorbed directly
RS domain A protein domain rich in arginine and serine from the substrate of dead organic matter after digestion by
dipeptides. RS domains are common in splicing factors in extracellular enzymes.
multicellular organisms. saquinavir Protease inhibitor of human immunodeficiency virus 1
RTK See receptor tyrosine kinase. (HIV1).
RU486 A drug that blocks progesterone receptor function. It is sarcoid An unusual granulomatous disease that affects many
used in breast cancer, induction of labour, and elective pregnancy organs and tissues including lung, skin, spleen and lymph node.
termination. Its cause is unknown.
Rubisco The enzyme ribulose 1,5-bisphosphate carboxylase/oxy- sarcolemma The plasma membrane of a muscle fibre.
genase. The carboxylase function is responsible for the fixation of SASP See small acid-soluble spore proteins.
carbon dioxide to ribulose 1,5-bisphosphate in the first step of the satellite A visible block of heterochromatin above the NOR region
C3 pathway. The oxygenase function is involved in photorespira- at the end of the short arm of acrocentric human chromosomes.
tion. satellite DNA Localized DNA sequences consisting of large
RuBP See ribulose 1,5-bisphosphate. numbers of tandem repeats, which often have base compositions
ruffle See lamellipodia. differing from the genomic average, and which therefore form
rule-of-six In the canine distemper virus, and similar viruses, the ‘satellite’ bands in caesium chloride equilibrium centrifugation
viral genomic RNA needs to be completely enwrapped by gradients.
nucleocapsid protein (N). A single N protein covers six satellite remote sensing Surveillance by satellite-borne sensors
nucleotides. To fulfil the rule, the number of nucleotides of the that has been used, e.g. to determine chlorophyll concentrations
genome must be divisible by 6. or water temperatures and can be used for tracking algal blooms
rumen The first vessel in the multichambered stomach of ruminant in water.
animals (including cattle, sheep, goats, deer, elk, bison, marsu- saturation (1) In genetics, describes the case where very large
pials, coloboid and langur monkeys, camels, hippopotamus, numbers of mutants of a sequence are made, so as to ensure that
llamas, and the hoatzin or stinkbird), in which cellulose is there is a mutant for any particular codon of the squence. (2) The
digested. case where two DNA sequences have such frequent multiple
run-through fraction In column chromatography, the portion of mutations at the same site that it is difficult to estimate an
a sample that flows unaffected by passage through a column accurate genetic distance between them. Such a situation is
made of some selectively adherent material. common among highly divergent sequences. (3) In regard to fatty
acids and similar lipids, the case where the hydrocarbon chain has
no double bonds, i.e. it contains the maximum possible number of
hydrogen atoms.
S chlorosomes Chlorosomes found in the anaerobic sulfur saturated fatty acid Fatty acid that contains no double bonds in
bacterial family Chlorobiaceae. the hydrocarbon tail, i.e. all the carbon atoms are ‘saturated’ with
S phase The DNA replication phase of the eukaryotic cell cycle. It hydrogen.
follows G1 and precedes G2 and mitosis (M phase). Saurischia The ‘reptile-hipped’ dinosaurs. These dinosaurs include
S0.5 The concentration of a substrate required to produce half- both herbivores and carnivores among their members and are
maximal activity. characterized by a more typically reptile-like arrangement of their
saccade A fast jerky angular displacement of the eye. hip bones.
Saccharomyces cerevisiae Baker’s or brewer’s yeast. Commonly sauropod dinosaurs Plant-eating dinosaurs with long neck and
referred to simply as yeast, this unicellular fungus is used as a tail, small heads, and five-toed feet. Examples include Bronto-
eukaryotic cell model system for scientific and industrial research. saurus, Diplodocus and Brachiosaurus.
saline agglutinin An antibody that directly agglutinates red blood Sauropodomorpha See sauropod dinosaurs.
cells suspended in saline. It is usually IgM. scaffold Protein around which a viral capsid is built.

232
Glossary

scale An organic or inorganic cell covering of variable shape that secretion leader sequence A sequence at the 50 end of a gene that
usually occurs in layers upon the plasma membrane of some types encodes a signal sequence in the protein which leads to the
of algae. targeting of that protein to the secretory system of the cell.
scaling The quantitative influence of body size on organic secretory granule Small intracellular vesicle that is packed with
function, including rate of metabolism. material and is destined for regulated exocytosis.
scanning the ATP-dependent, 50 to 30 movement of the 40S subunit sections, ultrathin Sections (less than 1-mm thick) prepared for
along a messenger RNA in search of an initiating AUG codon. electron microscopic resolution. Even bacterial cells must be
scansorial A type of locomotion characterized by rapid scurrying sectioned into more than 10 slices. Cells are solidified for
movements that is particularly common among small clawed microtomy with special knives made of diamond or glass.
mammals. Solidification is achieved either by deep freezing and slicing of
scattering The process whereby a wave is diffusely redirected in the still frozen block at low temperatures (cryosections), or
space. by stepwise replacement of cellular water (in bacteria about
SCE See standard calomel electrode. 80% w/w) by a synthetic resin.
SCF See stem cell factor. sedimentary rock A rock, such as sandstone or limestone, that was
Schiff base The C5N-containing covalent adduct formed by the once a sediment such as sand or mud, and which has not been
reaction of a primary amine with either an aldehyde or a ketone. modified substantially after burial.
schizogony An asexual multiple fission stage in the life cycle of segmenter Intracellular stage in which asexual division of
Apicomplexa (and other protozoa), leading to the formation of sporozoan protozoa occurs.
merozoites. The multinucleate cell divides into multiple progeny segmentation The division of the animal body or other structure
through simultaneous budding from the cell membrane into a series of segments of rather similar internal structure.
schizont Intracellular stage in which asexual division of sporozoan segregating site A nucleotide site at which two or more different
protozoa occurs. types of nucleotide are present in a population.
SCID See severe combined immune deficiency. selectins A family of cell adhesion proteins present on leucocytes
sclerenchyma Plant cell type characterized by the presence of and endothelial cells. They possess an extracellular lectin-like
lignified secondary cell walls and the absence of cellular contents domain that recognizes sugars on cell-surface glycoproteins.
at maturity. selection coefficient A measure of the strength of natural
sclerite A small scale, typically embedded in an epithelium. selection.
scleroderma A disease characterized by deposition of fibrous selection Non-random differential survival and reproduction (in a
tissue in skin and internal organs. particular environment) that is due to differences in genotype and
sclerotic cell Thick brown-walled globose to oval cells that thus phenotype.
reproduce by forming cross-walls and represent the tissue form
selectivity index Ratio of the TC50 to the IC50 (or, more broadly,
of fungi causing chromoblastomycosis.
toxicity to efficacy) for an antiviral drug. Should be as high as
scopuloid Ultrastructurally complex area in the suctorian swarmer possible, with a ratio of 100 often used as a benchmark.
where the suctorian stalk is secreted. It is not homologous to a
selenodont A condition found in some herbivores, in which the
‘scopula’ in other ciliates.
cusps of the molars are arranged in crescent-shaped crests.
score matrix For a dynamic programming algorithm, the matrix
selenoprotein A protein that contains the amino acid seleno-
that holds the cumulative scores between a query and the target
cysteine.
amino acid or nucleotide sequence.
scurvy A disease caused by vitamin C deficiency. SELEX Systematic evolution of ligands by exponential enrichment.
A technique that selects high-affinity nucleic acid sequences from
SDS-PAGE Sodium dodecyl sulfate–polyacrylamide gel electro-
a random pool.
phoresis. An electrophoretic technique that denatures and
separates proteins/peptides on the basis of their molecular mass. self The body’s own tissues.
SDV See silicon deposition vesicle. self assembly The ability of a complex macromolecular structure
second messenger Small diffusible intracellular signalling mole- such as a ribosome or virus particle to assemble itself from its
cule, produced as a result of receptor stimulation, that conveys components without external assistance.
the signal from the receptor to some enzyme or molecular system selfish DNA DNA sequences that are maintained solely by their
in the cell, which mediates the cell’s response. Common second capability to replicate within the genome, despite their capacity to
messengers are cyclic AMP, cyclic GMP, Ca21 , inositol 1,4,5- reduce the fitness of their hosts.
trisphosphate and 1,2-diacylglycerol. self-tolerance The normal ability of the immune system to
secondary chemical Molecular compound produced by an distinguish self from nonself, and to avoid mounting an immune
organism but not essential for its basic life functions. attack on self antigens.
secondary root See lateral root. semelparity A reproductive pattern in which the females normally
secondary structure Those regular elements of two-dimensional give birth only once in their lifetime.
structure common to many proteins that are formed by relatively semidiurnal tides A sequence of tides consisting of two high and
short-range interactions within the polypeptide chain. Examples two low tides over approximately 24 hours and 50 minutes. The
are a-helix and b-pleated sheet. high tides are of approximately equal amplitude, as are the low
secondary wood Woody tissue produced by lateral growth from tides.
the stele, and which provides the bulk of the stem in trees and semidominant mutation A mutation that results in a phenotypic
shrubs. effect when in the heterozygous state, but which is more severe in
secretory antigen Antigens secreted by an organism. the homozygous state.

233
Glossary

semi-persistent Describes the relationship between a plant virus serum sickness The clinical symptoms that accompany an allergic
and its arthropod vector in which the vector retains the virus for a reaction to the horse serum that formerly was administered to
period of hours to days. protect against diphtheria or tetanus.
senescence The steady decline in many aspects of adaptive seta (plural setae) A small hair-like projection formed from the
performance (e.g. sensory, cardiac, immunological) as a person insect cuticle.
ages during adult life. severe combined immune deficiency (SCID) Inherited immune
sensitivity amplification The production of a large percentage deficiency, usually rapidly fatal without treatment, in which both
change in the steady-state concentration of an active signalling T cells and B cells are very few or absent. It can be due to a variety
molecule in response to a small percentage change in a stimulus. of causes.
sensor In two-component bacterial sensory systems, a histidine severing Of actin, breaking of an actin microfilament.
protein kinase that can be autophosphorylated and then sex chromosome The chromosomes responsible for sex determi-
phosphorylate the response regulator as part of the signal nation (e.g. X and Y in humans) and which are present in different
transduction process. numbers or combinations in the different sexes.
sensory learning An improved capacity of the nervous system to sex determination The genetic, molecular and cellular mechan-
process the sensory information related to specific experiences, isms which direct development as a male or a female.
resulting in a more effective or efficient behaviour. sex differentiation Implementation of a sex-specific programme
sensory messages Signals recognized by sensory organs and of development.
which give information about physical and chemical environ- sex linkage The association of a genetic trait with one sex only,
mental characteristics (e.g. light, odorants, pheromones). which is usually due to the location of the gene for that trait on
sensory receptor A specialized nerve cell that introduces informa- one of the sex chromosomes.
tion into the nervous system by transducing specific events or sex pilus See F pilus.
conditions into a neural code.
sex-linked Describes a genetic trait associated with one sex only. In
sensory system A neuronal network made up of the sensory
mammals the clinically important sex-linked traits are usually
receptors and all successive processing centres of sensory
specified by genes on the X chromosome.
information in the nervous system. Sensory systems organize
sexual dimorphism The existence of a difference in structure and
the activity of large populations of neurons to make information
appearance between males and females of the same species. The
accessible to other brain systems.
term is sometimes also used to describe functional differences,
sensu stricto The most precise, restrictive or exclusive interpreta-
such as the frequency with which particular behaviours are
tion of a species (contrast with sensu lato).
displayed.
septum (plural septa) A cross wall formed within a fungal hypha
sexual reproduction Type of reproduction in which, at some stage
or at the base of a reproductive structure.
of the life cycle, haploid gametes (or haploid nuclei) are produced
sequence (1) A succession of layers of sedimentary rock. (2) The
as a result of meiosis. A gamete subsequently fuses with another
order of nucleotides in a nucleic acid or of amino acids in a
gamete of different ‘sex’ or mating-type (often from a different
protein.
individual) to produce a diploid cell (or nucleus) from which a
sequence similarity Two amino-acid sequences are said to show
new reproductive individual will develop.
similarity when there is a minimum of 30% identical amino acids
SF1 See splicing factor 1.
in comparable sequence regions.
shape The geometry of an organism after information about scale,
sequence-specific DNA-binding proteins Proteins that bind
with high affinity to sites on DNA consisting of a particular position and orientation has been removed.
sequence of base pairs. Sharpey fibres Extrinsic collagen fibre bundles by which bones
sequence-tagged site (STS) A short, known DNA sequence and teeth attach to muscles, tendons and ligaments.
which can be detected using the polymerase chain reaction. shear Deformation of a material caused by coplanar but noncol-
sequestration (1) Withdrawal of the more mature stages of the linear forces.
malaria parasite Plasmodium from the peripheral circulation shift-up An experiment in which cells growing slowly are suddenly
and their adherence to endothelial cells lining the deep tissues. shifted to a medium that allows more rapid growth.
(2) Keeping a reserve of some molecule in a particular form shoot apex The tip of a plant shoot, defined as the summit of the
or location. apical meristem. This is the most central point of the meristem
serine proteases A family of proteolytic enzymes that contain the and is often the highest point of the meristem where undiffer-
active-site triad of serine, aspartic acid and histidine residues in entiated meristematic cells reside.
their catalytic site. They include trypsin, chymotrypsin and shoot apical meristem The meristem located at the apex or
elastase. growth tip of the shoot.
seroconversion The appearance of antibodies in the serum after shoulder dystocia Difficulty in delivering the shoulder encoun-
exposure to an antigen. tered with abnormally broad-shouldered, big babies. Birth injury
serogroup A subdivision of a species (especially of bacteria) based may result.
on differences in surface antigens. Shungura Formation Block of sediments that lie to the west of the
serotype A subdivision of a serogroup, e.g. Leptospira interrogans Omo River just before it drains into Lake Turkana.
serogroup icterohaemorrhagiae serotype icterohaemorrhagiae. siamois A homeobox gene in Xenopus that encodes a likely
serovar See serotype. transcription factor which plays a role in inducing the Spemann
serum amyloid protein (SAP) A protein of the pentraxin family organizer.
which is an acute-phase protein in mice and is found in sick euthyroid syndrome Abnormal thyroid function blood tests
amyloidosis. in the absence of thyroid disease.

234
Glossary

side chain The group of atoms that are connected to the a-carbon Sjögren syndrome A syndrome involving dryness of mucous
of an amino acid. membranes and inflamed parotid glands.
siderite An iron carbonate mineral, FeCO3. skeleton The supporting hard tissues in an organism, either
siderophore A chelating protein, produced by both plants and internal or external.
microorganisms, with a high affinity for metal ions. skewness Pertaining to the asymmetry of a frequency function as a
sigma factor A subunit of bacterial RNA polymerase that is departure from the normal distribution.
required for initiation of transcription. Some sigma factors confer skull The bony casing of the brain (the cranium) and the mandible.
the ability to recognize and bind to a particular promoter, thus skull roof A series of dermal bones of the skull that protect the
changing the pattern of gene expression. braincase from above, including (in turtles) the nasal, prefrontal,
signal peptidase A protease that specifically cleaves the signal frontal, parietal, jugal, quadratojugal, squamosal and postorbital
peptide from the N terminus of a protein. bones.
signal peptide A stretch of amino acids, located at the N terminus S-layer Self-assembled paracrystalline surface arrays consisting of
of a protein, that targets it to be secreted. protein or glycoprotein which reside on top of the cell wall of
signal transducers and activators of transcription (STATs) A prokaryotes.
family of transcription factors that are defined by sequence sleeping sickness Disease caused by Trypanosoma brucei gam-
homology and are considered to be the primary substrates of the biense or T. b. rhodesiense, restricted to sub-Saharan Africa.
receptor-linked Janus kinases. slime Polysaccharide- and/or glycoprotein-rich material secreted
signal transduction A series of interlinked biochemical reactions by many types of organism. In bacteria, it is similar to the
in which a signal initiated at the cell surface is transmitted inside capsular material but is not attached to the cell wall.
the cell to cause a specific cellular response. slimy Describes a fluid with a low degree of shear thinning. A slimy
silicon deposition vesicle (SDV) A special endomembrane food material is one that coats and does not clear the mouth; it
component of golden algae where silica-complexed coverings may also be difficult to swallow.
are produced within the cell. slit sensillum (plural slit sensilla) A sense organ unique to
similarity search A search for sequences similar to a query arachnids that appears as a small slit in the cuticle. It is sensitive to
sequence in the DNA and amino-acid sequence databases by compressional forces in the exoskeleton that act perpendicular to
pairwise comparison of sequences using alignment tools. the long axis of the slit.
simple spike Action potential with a simple configuration, induced small acid-soluble spore proteins (SASP) Low-molecular weight
in a Purkinje cell by activation via parallel fibres. proteins produced during bacterial endospore formation that
single-strand annealing A DNA repair process involving diges- bind to DNA to protect it against damage and serve as a source of
tion of the opposing single strands of a DNA duplex on either side metabolizable amino acids for germinating spores.
of a double-strand break to expose regions of homology on the small nuclear ribonucleoprotein particle (snRNP) A stable
remaining strands. These regions can then pair to facilitate repair. complex of small nuclear RNAs with proteins. Such particles are
single-stranded DNA-binding protein Protein that binds non- involved in the RNA splicing reaction in the nucleus.
specifically to single-stranded DNA with much higher affinity small nuclear RNA (snRNA) Small RNA molecules present in
than to either RNA or double-stranded DNA. the nucleus usually complexed with protein to form small
sink tissues Tissues dependent upon the inward flow of nutrients nuclear ribonucleoproteins. The snRNAs involved in
for growth. pre-mRNA splicing are U1, U2, U4, U5 and U6, and, for the
sink A region of a plant in which the net flux of solutes into sieve splicing of U12-dependent introns, U11, U12, U5, U4atac and
elements is sufficient to produce net export from them. U6atac.
sinter Porous silica-rich deposit of hot springs and geysers. It forms smectite Clay mineral with a very high proportion of the mineral
terraces and mounds in hot-spring areas. montmorillonite.
sister chromatids The two copies of a chromosome after its smoker A hot deep-sea vent with precipitate-forming plume
replication and while they are still attached to each other. (‘smoke’). The precipitates may be black or white (‘black
sister group In phylogenetic systematics (cladistics), either of two smokers’ and ‘white smokers’).
species groups that arose from the stem species of a monophyletic smooth muscle Type of muscle found e.g. in blood vessel walls, gut
group in the same splitting (speciation) event. Sister groups have walls and in other organs that are under involuntary control by
the same rank. Furthermore, any newly arising species has a sister the nervous system. When looked at under a microscope, the
group by definition. small cells making up this type of muscle have no crossbands or
sister taxa Taxa sharing a more recent common ancestor with each striations.
other than with any other known taxa. SN2-reaction Nucleophilic substitution. The first and rate-limit-
sister union (SU) A type of deletion in sister chromatids in which ing step of the reaction is a collision of the original complex with
the remaining sister chromatids become united at the break point, the nucleophile. The reaction is therefore bimolecular and of the
and the deleted acentric portions also become fused at the broken second order.
ends. SNC meteorites A group of meteorites originating from the same
site-directed mutagenesis The production of defined mutations parent body, probably Mars.
at defined positions in a DNA sequence. snRNA See small nuclear RNA.
site-specific recombination Process in which specific segments of snRNP See small nuclear ribonucleoprotein particle.
DNA are synapsed by site-specific recombinase proteins which social behaviour A behaviour that affects not only the fitness of
then catalyse conservative breakage–rejoining reactions to the individual performing it, but also the fitnesses of nearby
generate recombinant DNA. organisms of the same species.

235
Glossary

sodium channel A voltage-sensitive transmembrane ion channel special pair chlorophyll A pair of chlorophyll molecules that are
present in the membranes of most neurons. When this channel is arranged in the reaction centre complex to cause charge
opened by a depolarizing voltage, sodium ions selectively pass separation.
through. specialist An organism with a restricted realized niche.
sodium dodecyl sulfate–polyacrylamide gel electrophoresis speciation Process in which new species are generated either by
See SDS-PAGE. splitting (cladogenesis) of an ancestral species or the gradual
soft selection Selection based on the relative fitness (rank) of transformation of one species into another (anagenesis).
individuals in a population. It is usually frequency dependent. species The most commonly used unit of biodiversity, defined as a
solfatara A hot, sulfur-rich, often acidic environment. The name is group of organisms of the same type that can interbreed with each
derived from the Solfatara Crater near Naples (Italy). other. A species thus comprises all the populations that occur
solid-phase immunoassay Type of immunoassay, such as across its overall range.
ELISA, in which one of the reactants (antigen or antibody) is specific activity The catalytic activity of an enzyme per milligram
linked to a solid support. protein.
Solnhofen limestone Sequence of Late Jurassic calcareous specific metabolic rate Metabolic rate per unit body mass, which
deposits famed for the excellent preservation of many different is useful in comparing metabolism in animals of different sizes.
types of fossils including pterosaurs and Archaeopteryx. specification Processes by which cell fate is acquired during
solvation Transfer of solute from the gas phase to solution. development.
solvent transfer The solvent transfer free energy of a molecule is a specificity In regard to symbiosis, the taxonomic range of a partner
measure of the propensity of the molecule to dissolve in a with which an organism forms a symbiosis. The specificity may be
nonaqueous liquid such as cyclohexane rather than in water. narrow (the organism associates with few, closely related taxa) to
soma (1) The animal body, with the exception of the cells of the broad (the organism associates with many, taxonomically diverse
germline. (2) The neuronal cell body containing the cell nucleus. taxa).
somatic cell Any of the cells that make up the body of an animal, spectrum Absorbance of a substance measured as a function of the
with the exception of the germ cells. wavelength or the frequency of the incident light.
somatic fate All cell fates other than the germline. Spemann organizer A signalling centre above the dorsal lip of the
somatic hypermutation An accumulation of point mutations blastopore in the Xenopus embryo which is responsible for
that occurs in the variable regions of rearranged immunoglobulin induction of the central nervous system and other tissues of the
genes in activated B cells. It results in a further diversification of dorsal axis. When transplanted to the ventral side of a host
antibody specificity and binding strength for the antigen. embryo, it can induce a complete new embryonic axis. Named
somatic mutation Mutations that occur in somatic cells, not germ after its discoverer, the Nobel prize-winning developmental
cells, and thus are not passed on to the next generation. biologist Hans Spemann.
somatic Describes a cell or structure that does not produce spermatogenesis The formation of sperm.
reproductive structures or gametes. spermatogonia (singular spermatogonium) The primitive
sonic hedgehog A gene related to the hedgehog gene, first germ cells that undergo meiosis to produce spermatozoa.
characterized as encoding a secreted protein signal in Drosophila. spermatophore A sperm package, consolidated by protein or
It has been found in many mammalian species, including humans, polysaccharide and transferred from male to female in several
and plays a critical role in patterning of vertebrate embryonic animal groups. Use of spermatophores circumvents copulation.
tissues. spheroplast A bacterial or yeast cell from which the bulk of the cell
sorocarp A fruiting body produced by cellular slime moulds in wall has been removed enzymatically under osmotic protection.
which the spores are not contained within a common wall but are spherule A sac-like structure producing asexual spores endogen-
instead suspended in a matrix of slime. ously that is the tissue phase of the fungus Coccidioides immitis.
sorus (plural sori) The mass of cells within a sorocarp. spherulocyte A type of haemocyte characterized by prominent
SOS mutagenesis An error-prone DNA repair mechanism in membrane-bounded, electron-dense, intracytoplasmic spherules,
bacteria that occurs under conditions of extreme DNA damage. which are considerably larger than granules in the granulocytes.
source A region of a plant in which there is net efflux of solutes and spicule A sclerotized intromittent organ that the male nematode
water from sieve elements, resulting in import into them. employs to hold open the female vulva during the transfer of
source tissues Tissues capable of assimilating carbon by photo- sperm at mating.
synthesis and exporting it to sink tissues. spine A protrusion of a postsynaptic dendrite that makes synaptic
Southern blotting A technique used for detection of a specific contact with a presynaptic cell’s axon terminal.
restriction fragment against a background of many other spinneret An appendage or projection in arachnids specialized for
restriction fragments. producing and handling silk. Found on the opisthosoma of
spasticity Muscle tightness and stiffness resulting from injury to spiders and the chelicerae of pseudoscorpions.
the brain or spinal cord. spiral cleavage Pattern of zygote cleavage found in species of
spatiotemporal pattern The information contained within a annelids, molluscs, sipunculans, entoprocts, platyhelminths and
group of neurons, which is represented both by the time-varying nemertines. It is characterized by alternation in the direction of
activity of individual neurons and by the particular set of neurons the mitotic spindles such that blastomeres are displaced so that
that becomes active. they lie in the furrows between one another.
spawn Substrate that has been impregnated by mushroom spiral valve Intestine with a longitudinal secondary wall laid down
mycelium and is used for mushroom production. The genetic as a spiral around a central axis.
and cultural characteristics of the mushroom species are carried spiralians Animals with a spiral type of cleavage, e.g. flatworms,
in the spawn. ribbon worms, molluscs, annelids.

236
Glossary

Spitzenkörper A cluster of vesicles located just inside the fungal stabilizing selection Selection acting against individuals at both
hyphal tip that is believed to be involved in the transport of wall extremes of a phenotypic distribution (e.g. selection against both
material in a polarized fashion. very small and very large individuals).
splanchnomegaly See organomegaly. stable isotopes Isotopes that do not undergo radioactive decay.
spliceosome A large complex of RNAs and proteins responsible standard apparent reduction potential E0 1 is the standard
for pre-mRNA splicing. A spliceosome forms around each intron reduction potential at a specified pH. It is defined by: E0 5
and dissociates after splicing is complete. E0 1 – (ln Q0 (RT/nF)).
splice-site mutation A mutation in a consensus DNA sequence standard calomel electrode (SCE) A widely used reference for
that is essential for effective RNA splicing. Splice-site mutations establishing redox potential. Redox potentials vs SCE 5
often result in the loss of exons from the messenger RNA (exon potentials vs SHE (standard hydrogen electrode) 1 0.242 V (at
skipping). 251C). Redox potentials vary with temperature and pH.
splicing factor 1 (SF1) A protein that interacts with the pre- standard rate of metabolism The rate of metabolism in adult
mRNA branchpoint during RNA splicing. Also known as BBP ectotherms at rest, at a particular body temperature, and when
(branch site-binding protein). post-absorptive.
spongiform changes Formation of sponge-like vacuoles within stapes A small stirrup-shaped bone in the mammalian middle ear.
the grey matter of the brain as a characteristic morphological It transmits sound vibrations to the inner ear, where they are
correlate in all prion diseases. transduced into electrical impulses.
spongiotrophoblast The outer layers of the placenta in the start codon The trinucleotide sequence in messenger RNA at
rodent. which protein synthesis starts.
spontaneous cytotoxicity See natural cytotoxicity. starter and extender units Materials that initiate formation
sporangiole A very small sporangium that contains only one or (starter) and elongation (extender) of carbon chains. In general,
two sporangiospores, and also lacks a columella. 1 starter unit + n extender units gives n CO2 + product.
sporangiophore A modified hypha that supports the sporangium. statocyst Gravity-detecting sense organ.
sporangiospore An asexual spore formed within a sporangium.
status migrainosus Daily and continuous headaches associated
sporangium (plural sporangia) The mother cell from which
with psychogenic tension or analgesic or other drug misuse.
asexual spores are formed, and which eventually develops into a
stavudine Nucleoside analogue inhibitor of human immunodefi-
sac-like structure containing the asexual spores.
ciency virus (HIV) reverse transcriptase. Used as an anti-HIV
spore A general term for a propagule that is formed either
drug.
asexually or sexually, is usually surrounded by a rigid cell wall,
steady state (1) A state of a macromolecule–ligand system in
and which is the dissemination stage of the life cycle.
which the equilibrium condition dC/dt is pertinent for only a part
sporidium Sexually differentiated basidiospore.
of the reaction components and which is temporarily achieved in
sporoblast A cell that develops directly into a spore without
a finite time interval. (2) A kinetic phase during enzyme catalysis
further division.
in which the rate of formation of the enzyme–substrate (ES)
sporocarp A multicellular fruiting body in which the spores are
complex is exactly matched by its rate of consumption. Hence,
contained by a common wall. Sporocarps are produced by the
during steady state, the concentration of ES complex is constant.
plasmodial slime moulds.
(3) For a gas-exchange system, a state in which the total masses,
sporogenesis See sporulation.
concentrations, partial pressures and fluxes of a given moving gas
sporogony (1) Mitotic fission stage in the life cycle of apicomplex-
species remain the same with time in all locations, e.g. in which the
an protozoans, leading to the formation of spores. (2) Develop-
amounts of gas consumed (oxygen) or produced (carbon dioxide)
ment of the fertilized zygote through a process of meiotic and
in the tissues equal those exchanged with the ambient medium.
mitotic divisions to form infective sporozoites. (3) In microspor-
steatosis Accumulations of fatty droplets in a tissue.
idians, the process of division of the sporont to form a sporoblast.
sporomorph Plant spore or spore-like object. stele The vascular tissue and associated pith in a stem or root,
sporont In microsporidians, a cell that is committed to sporogony which translocates substances from one part of a plant to another.
and, after a specific number of divisions, becomes sporoblasts. stellate Star-shaped.
Sporonts are sometimes characterized by a membrane-dense stem cell An undifferentiated cell that is capable of continued
ribosome-rich cytoplasm and a thickening of the cell membrane. indefinite division, giving rise both to more stem cells and to other
sporophyte Phase of plant life cycle that forms spores. daughter cells that undergo differentiation into particular cell
sporotrichosis Invasive fungal infection caused by Sporothrix types.
schenckii. stem cell factor (SCF) A protein required for the in vivo growth of
sporulation The formation of spores and their associated specia- haematopoietic stem cells. In vitro it stimulates the growth of blast
lized supporting and containing structures. cell, erythroid and granulocyte colonies.
SpoT An enzyme with (p)ppGpp 30 pyrophosphohydrolase activity stem lineage If a species becomes the stem species of a clade
and potential (p)ppGpp synthetic activity, closely involved with through splitting, its lineage becomes the stem lineage of the
RelA in the ‘stringent response’ of bacteria. clade.
squamates A group of reptiles (the Order Squamata) consisting of stem–loop An RNA structure formed by pairing of self-comple-
snakes, lizards, and amphisbaenians. They are characterized by mentary regions flanking an unpaired loop sequence.
reduction or absence of limbs, elongated bodies and tails, stenothermal Tolerant of only a narrow range of temperature.
keratinous scales, vomeronasal organs and paired copulatory stenotopic Having a narrow environmental range.
organs called hemipenises. stepping-stone model migration The movement of individuals
SSV1 Temperate virus of the archaeon Sulfolobus shibatae. among neighbouring populations.

237
Glossary

stepwise mutation model A model for the formation of tandem stratigraphic debt The sum of range extensions measured in
arrays (e.g. a microsatellite array) which proposes an increase or temporal or stratigraphic units.
decrease by one repeat unit at a time. stratigraphy The arrangement of rocks in sequence in the Earth’s
stereocilia (singular stereocilium) Modified microvilli of the crust. The study of the dating and sequence of rocks.
hair cells of the cochlea that serve as mechanoelectric transducers. stratocladistics An amended version of cladistics in which both
stereoelectronic effects Molecular conformation-dependent implied stratographic gaps and implied homoplasies are con-
influences on reaction rates due to the geometric dependence sidered evidence against a possible phylogeny.
of atomic orbital interactions. strepsirhine A cladistic term of primate classification that includes
stereotyped song Adult stable form of a bird’s song. only the lemurs and lorises.
sterigma (plural sterigmata) A spike-like protrusion on a stress (1) Physiologically defined in humans and other mammals as
basidium that supports a basidiospore. a response of the organism to changes in its homeostasis,
sterile cabinet An enclosure that provides an aseptic work area for specifically involving activation of the hypothalamic-pituitary-
culture. adrenocortical (HPA) axis, resulting in the secretion of
sterile Tissues or structures that are not producing spores. corticotrophin-releasing factor, adrenocorticotrophic hormone
sterilization The killing of all living organisms on or in a material. (ACTH) and corticosteroids, and in coactivation of the
Sternberg–Reed cells Large neoplastic binucleate cells character- sympathetic nervous system and the adrenal medulla. (2) The
istic of Hodgkin disease. force in a structure acting over an area that resists internal forces.
sternite A cuticular plate that occurs on the ventral surface of each stress fibres Bundles of relatively stable actin filaments spanning
body segment in arthropods. the length of cells, involved in maintaining cell shape.
sterol A hydroxylated tetracyclic compound that retains some or stress-induced premature senescence Long-term appearance
all of the carbon atoms of squalene in its nucleus and side chain of the biomarkers of replicative senescence after exposure of
and partitions nearly completely into the ether layer when it is human diploid cells to sublethal stress.
shaken with equal volumes of water and ether. striated muscle The skeletal muscle that forms the main mass of
stewardship The responsibility to protect biological diversity, as the limbs and much of the body and is responsible for the ability
given by God or accepted from society. to move. It is generally under conscious or voluntary control.
sticking The stationary arrest of blood cells in the presence of fluid When looked at under a microscope, the long cells making up this
shear, mediated by secondary adhesion receptors. type of muscle have crossbands or striations.
stigmata of congenital syphilis The signs that are specific or stroke volume The volume of blood pumped by one ventricle
common in congenital syphilis: saddle nose, interstitial keratitis, during a single heartbeat.
fibrosis around angles of the mouth, Hutchinson teeth or Moon stroma Liquid phase of the chloroplast, located outside the
molars, palatal gumma, Clutton joints. thylakoid membrane vesicle, where photosynthetic carbon
stochastic Random, by chance. reduction, starch synthesis and other important biosynthetic
stoichiometric Present in equimolar amounts. pathways take place.
stomata (singular stoma) The pores in the underside of a leaf stromal cell The supportive tissue of an organ.
through which gas exchange takes place. Each is composed of a stromal membranes Unstacked thylakoid membranes that
pair of specialized cells whose movement opens or closes the pore. extend between grana stacks in chloroplasts.
stomatogastric nervous system A part of the nervous system stromatactis A common feature of many Palaeozoic reefs of
that supplies and controls the anterior part of the gut in multiple and controversial origin, consisting of cement-filled
arthropods. laminar voids with flat or undulose lower surfaces and irregular
stomium A cluster of cells that control the rupture or dehiscence of or digitate upper surfaces.
pollen grains. stromatolite A laminated calcareous structure produced by the
stomodaeum An intucking of ectoderm meeting endoderm in activities of lime-secreting cyanobacteria.
Ctenophora, corresponding to the mouth of higher invertebrates. strongyles stylet A relatively solid, sclerotized structure in the
stop codon The trinucleotide sequence in messenger RNA at mouth area of plant-parasitic nematodes. It is used to transfer sap
which protein synthesis terminates. from plant cells to the nematode pharynx.
stopped-flow spectroscopy Method of rapid mixing of two fluids structural imaging The formation of an image of anatomical
that is used to capture fast reactions. The fluid flow is stopped by a structure.
mechanical mechanism, and the mixture is observed immediately. structure factor The X-ray scattering along a particular direction
strabismus Commonly known as squinting, an abnormal devia- from a single unit cell, equivalent to the diffraction pattern sample
tion of an eyeball preventing a binocular focus of the same visual only at reflection peaks.
target. STS See sequence-tagged site.
strain energy Ability of matter to do work by virtue of internal stylet In insects and mites, mouthparts modified into a piercing–
deformation. sucking instrument. In plant-parasitic nematodes, a single
strand break A type of DNA damage in which a break occurs in stiletto-like part of the foregut (odontostyl) needed to open
the sugar–phosphate backbone of DNA. A break on one side of epidermal cells or even to reach deeper tissue layers.
the helix is called a single-stranded break, while matching breaks stylet bundle The feeding organ of the Hemiptera (bugs). It is
on both sides of the helix are called double-stranded breaks. composed of two external mandibular stylets and two maxillar
stratification The difference in density at different depths in a stylets. Longitudinal grooves in the maxillar stylets form the wide
column of water, which is dependent on variation in temperature, food canal for ingestion and the narrow salivary canal for
pressure and (in seawater) salinity. egestion.

238
Glossary

stylet-borne transmission Common type of transmission for superantigens Proteins that stimulate large numbers of T cells
plant diseases with aphid vectors. Acquisition and infection in an antigen-nonspecific fashion. They simultaneously bind
feeding times are short, there is no lag-phase, the vector is certain types of T-cell receptors and MHC class II molecules (at a
infective for short times and infectivity is lost on moulting. site distinct from the regular antigen-binding site).
subacute sclerosing panencephalitis A rare disease caused supercoiled Describes a closed DNA double helix which is itself
by persistent measles virus infection in the central nervous coiled around its own axis.
system. supercoiled DNA Closed circular DNA formed from linear DNA
subcellular marker A selected biomolecule that is confined to a after twisting (positive) or untwisting (negative) the DNA helix.
specific organelle or suborganellar compartment and that may supercoiling Coiling superimposed on that of the helix of the DNA
be used to determine the presence of this compartment in a molecule itself.
subcellular fraction. supercooling point Temperature below the freezing point to
subduction The process in which one crustal plate plunges beneath which a solution can be cooled before spontaneously freezing.
another, eventually reaching the mantle and melting. superficial Describes a noninvasive cutaneous or subcutaneous
suberin Aliphatic biopolymer composed of linear, long-chain, infection.
lipophilic compounds derived from elongated fatty acids. superinfection immunity The inability of a bacteriophage to
subgenomic RNA A viral messenger RNA that is identical in infect a bacterial cell lysogenic for the same phage. This is due to
sequence to the 30 -end of the genomic RNA. the turn-off of genes on the infecting genome by the resident
subsensitive response In an individual level of regulatory prophage repressor.
cascade (such as an intracellular signalling pathway), an superior Towards the head.
amplification response that is more gradual than a Michaelian
supermolecule A discrete structure comprised of more than one
response.
compound, usually polymers.
subspecies Fully interfertile geographic races of the same species
supernova An explosion of a massive star which ejects most of the
that have been assigned formal taxonomic names.
original mass into space.
substituted galacturonans A pectic polysaccharide composed of
superoxide anion A derivative of oxygen capable of oxidative
a 1,4-linked galacturonic acid backbone that is substituted at C2
destruction of cellular components.
and/or C3 with mono- or oligosaccharide side chains.
superoxide dismutase Enzyme that converts superoxide radicals
substitution (1) A mutation that replaces one nucleotide with
formed after photosynthetic reduction of oxygen to hydrogen
another in a DNA molecule, or results in the substitution of one
peroxide (H2O2).
amino acid with another in a protein molecule. (2) Replacement
of one allele by another in a population. suppressor mutation A secondary mutation that totally or
substitution matrix A matrix containing scores for weighting the partially restores a function lost as a result of a primary mutation,
various possible substitutions between different characters, most and which is located at a site different from that of the primary
commonly amino-acid residues. mutation.
substrate (1) The molecule(s) of reactant that are acted upon by suppressor of PEV (Su(var)) A second-site mutation that leads to
an enzyme and thus converted to the products of the reaction. suppression of the position-effect variegation (PEV) phenotype,
(2) The surface upon which an animal is positioned or moving, i.e. one that results in more expression (less silencing) of a
such as the ground, a tree branch, etc. euchromatic gene subject to PEV.
substrate specificity The property of an enzyme that it will act suppressor T cells T lymphocytes that negatively regulate the
only (or most efficiently) on one substrate. extent of an immune response, and can even promote nonrespon-
substrate-level phosphorylation Process in which the generated siveness or tolerance to an antigen.
phosphoryl group is formed attached to the actual substrate of an supraclavicular In the hollow above and behind the clavicle.
enzyme. supramolecular structure An aggregate of a number of individual
substratum (plural substrata) Organic matter that serves as a molecules held together by noncovalent forces.
food source and support for fungal growth. supraorbital ridges Brow ridges above the eye orbits that are
subtilisin A serine protease produced by sporulating cells of present in some archaic hominid species. In Homo erectus these
Bacillus subtilis and which is used in washing powders. form a continuous shelf of bone, but in the Neanderthals they are
subunit vaccine A vaccine that contains only one or a few curved over each orbit and do not connect in the midline.
individual purified components from the microorganism it is supravalvular aortic stenosis (SVAS) A narrowing of the artery
designed to protect against. that is above the valve of the heart’s aorta.
sugar mimic A substance that, although not a sugar, has a similar surface coil A radio transmitter/receiver that transmits radio-
molecular shape and can therefore lodge at the active site of an frequency energy to, or receives signal from, tissue that is
enzyme. This prevents the sugar docking with the enzyme and immediately adjacent to the plane of the coil. Such coils are
thus prevents metabolism of the sugar. typically constructed as one- or two-turn solenoids.
SU See sister union. surrogate A person appointed to make medical decisions for an
sulfate reduction Anaerobic bacterial metabolism that utilizes individual who is no longer able to make those decisions for
sulfate (SO42) dissolved in sea water in the metabolism of themself.
organic matter, and produces hydrogen sulfide (H2S) and suspension feeding Mode of feeding by removal of suspended
bicarbonate (HCO3) as byproducts. food particles from the surrounding medium.
sulfur globules Deposits of organosulfanes present in the peri- suspensor A hyphal tip that at first supports the gametangium and
plasmic region of certain bacteria. then the zygospore.

239
Glossary

sustainability The ability of a society or a particular human synapsis (1) The bringing together of two transposon ends by
activity to continue indefinitely without depleting resources or transposase–transposase interactions. (2) The pairing of two
damaging the environment. homologous chromosomes during meiosis.
SVAS See supravalvular aortic stenosis. synaptic cleft The extracellular-matrix-filled gap separating
swarm cells Uninucleated cells with flagella which emerge from individual neurons at a synapse.
amoebae in the life cycle of Physarum. synaptic depression Reduction in synaptic transmission, usually
switch I Structural element in G proteins which changes con- due to depletion of a presynaptic store of transmitter.
formation depending on whether GDP or GTP is bound. synaptic eligibility trace A transient neurochemical event
switch II Structural element in G proteins which changes orienta- that serves as a memory of recent synaptic activity, and
tion depending on whether GDP or GTP is bound. indicates that a particular synapse is ‘eligible’ for adaptive
symbionin A GroE-like protein synthesized by the endosymbiotic modification if an error in performance is subsequently
bacteria of aphid cells. detected.
symbiont Most generally, a species partner in a symbiosis. Also synaptic plasticity The amount of neurotransmitter released at
frequently used, in contrast to host, to refer to the smaller member synapses of many neurons can vary with the past history of the
of the association. synapses. Changes in synaptic strength are thought to be the basis
symbiosis An intimate association between two species, which may of learning and memory.
be advantageous to both partners (mutualism), be of advantage synaptic potential A brief, graded voltage change in a neuron
to one partner at the expense of the other (antagonism), or be generated by the opening of ion channels following the release of
neither harmful nor advantageous to either (commensalism). neurotransmitter from the presynaptic terminal.
symbiosome A membrane-bounded vacuole containing a sym- synaptic strength This is measured by determining the average
biont(s) within the host tissue. amplitude of the excitatory postsynaptic response caused by a
sympatric Occupying the same geographical location. presynaptic action potential.
sympatric speciation Formation of a new species without synaptic vesicles Small membrane-bounded sacs inside the axon
geographical separation from ancestral species. It is considered terminal that contain chemical neurotransmitters (e.g. acetylcho-
uncommon, hence the significance of polyploid speciation as an line) that are released from the axon by exocytosis in response to
example. the arrival of an action potential.
sympatry Occurring in the same geographic region, thus having synaptosomes Isolated nerve endings in homogenates of neural
the potential to interbreed. tissue formed when synapses break off from associated axons and
symplasm The interconnected cell contents of a plant. their plasma membrane reseals.
symplast The interconnected cytoplasms of a plant, as a whole. syncope Sudden loss of consciousness.
symplastic Describes the supracellular organization of the inter- syncytium (1) A multinucleated cell formed by the cytoplasmic
connected cytoplasm in plant tissues. fusion of individual cells. (2) A population of cells whose
symporter A transmembrane protein that couples the transport of cytoplasms are connected by gap junctions is also sometimes
two solutes across a membrane in the same direction. referred to as a syncytium or a syncytial network.
symptomatic infection An infection that produces clinical signs, syndrome A collection of symptoms and features associated with a
based on which the infection can be diagnosed. specific disease.
symptomatic migraine Migraine-like attacks that are rarely syneresis The expulsion of aqueous solution from a gel caused by
caused by brain diseases, such as tumours, malformations, raised shrinking of the gel. Characterized by the formation of droplets
intracranial pressure, strokes and inflammatory conditions of the or a liquid layer over the surface of the gel.
blood vessels. synergism See synergy.
synaptic facilitation See facilitation. synergistic Acting cooperatively to have an enhanced effect, e.g.
synanthropic Typical of human dwellings and benefiting from, or two mutations which individually have only a mild or no
able to exploit, human activities. phenotype, but when combined are lethal.
synapomorphic Describes a derived or specialized homologous synergy, synergism The case when two agents or stimuli acting
character found in species of two or more groups that is together have a greater effect than either acting alone.
hypothesized to have arisen in a directly ancestral group and syngeneic graft See isograft.
not in any earlier ancestor. The term is popularly used in cladistic syngeneic mouse strains Strains of mice possessing almost
studies. identical genotypes.
synapomorphy A shared derived character. In cladistics, a synteny The occurrence of a given set of genes in the same order on
character that defines a clade. a chromosome in different species.
synapse The point of near-opposition of one neuron with another synthetases Enzymes that catalyse the hydrolysis of ATP to ADP
at which signals are passed from one cell to the other. Usually, the and PI coupled to the formation of X-Y from X and Y.
end of the axon of one neuron forms a synapse on a dendrite of syntrophy Microbial interaction in which two species working
another. Neurotransmitter is released from the cell on one side of together carry out a particular chemical transformation that
the synapse (the presynaptic side) and triggers an electrical charge neither organism could perform alone. Literally means ‘eating
in the cell on the other (the postsynaptic side). together’.
synapse specificity The phenomenon in neuronal plasticity that systematics (1) Study of the diversity of organisms, their
not all synapses on the same neuron have to change together. classification and nomenclature. (2) The process of organizing
Depending on stimulation conditions, some synapses can get taxa (species and higher categories) into a hierarchical pattern
stronger, others can get weaker, and many may not change at all. that may approximate their evolutionary relationships.

240
Glossary

systemic acquired resistance In plants, a resistance response taphonomy The study of the processes of fossilization, essentially
manifest throughout the whole plant that results from either those processes that act on a once-living organism following
pathogen infection or chemical treatments, and often confers a death. It includes those that act upon a carcass between death and
durable and broad-spectrum disease resistance. final burial, and the changes in the chemistry of the fossil that
systemic circulation See peripheral circulation. mainly occur after burial. It may yield forensic evidence
systemic lupus erythematosus An autoimmune inflammatory concerning the life habits of the fossilized organism and the
connective tissue disease with variable features frequently circumstances surrounding the death.
including fever, fatigue, joint pains, and diffuse reddening of tautomers Chemical isomers that differ only in the location of
the skin of the face and upper body. hydrogen atoms. They exist in rapid equilibrium, although one
systemic response Activation of wound-response genes in the form is usually more stable.
undamaged tissues of a wounded plant. The term usually refers to taxon (plural taxa) A group such as Mammalia or Homo sapiens
responses occurring in different leaves from the one actually that is a unit, rather than a kind of unit (e.g. family, class), in a
wounded. formal classification system.
systemin Peptide of 18 amino acid residues, derived from the N- taxonomy The science of identification, classification and nomen-
terminal part of a 200 amino acid precursor protein, apparently of clature of organisms.
cytosolic location. A potent inducer of proteinase inhibitor TC50 Drug concentration that is toxic to 50% of cells.
expression in tomato plants. T-cell antigen receptor See T-cell receptor.
systole The period of contraction of the ventricles of the heart. T-cell receptor (TCR) The antigen-specific receptor of T cells,
syn conformation Conformation of nucleotide where the encoded by genes that undergo rearrangement during T-cell
Watson–Crick face of the nucleobase is oriented over rather development. It is a cell-surface heterodimer of either a and b
than away from the sugar. chains (the majority of T cells) or g and d chains, complexed with
signal-transducing proteins. T-cell receptors recognize peptide
antigens complexed with MHC molecules, and each T cell bears
T cell-dependent antigen An antigen that requires the coopera- receptors of only one specificity.
tion of CD4 T cells with B lymphocytes for an effective antibody Tcf An HMG-box transcription factor that binds DNA with
response to be made. Most protein antigens are T-cell dependent. sequence specificity, altering the bending of the DNA and thus
T cells Thymus-derived lymphocytes, the lymphocytes that mature gene expression.
in the thymus and are the cells responsible for antigen-specific TCR See T-cell receptor.
cell-mediated immunity and which, in most cases initiate adaptive TDLU See terminal duct lobular unit.
immune responses. They include the CD4 (helper T cell) subset T-DNA Fragment of the Ti plasmid transferred to plant cells during
(which secretes cytokines acting on B cells, macrophages and Agrobacterium infection (‘transferred DNA’).
other T cells) and the CD8 subset, which includes cytotoxic T TdT See terminal deoxynucleotidyltransferase.
cells. tectonic Refers to processes or features of deformation of rocks,
T lymphocyte See T cells. commonly during mountain building or movement of continents.
T7-transcript RNA molecule synthesized in vitro by transcription tectum The dorsal part of the midbrain, which contains maps of
of a synthetic or natural gene by bacteriophage T7 RNA the space around an animal and is involved in multisensory
polymerase. integration.
tachyphylaxis Rapid loss of response. teichoic acid, teichuronic acids Polymers of glycerol and ribitol
tagmata See tagmosis. joined by phosphate groups, with amino acids or sugars attached.
tagmatization The regional specialization of groups of segments Major components of the Gram-type positive cell wall, these
in arthropods to form functional units or tagmata (e.g. head, polymers are found in great variety and are highly antigenic.
thorax, abdomen). teleomorph The sexual stage of a fungus, e.g. the cup in an
tagmosis Regional specialization of the body. ascomycete or the mushroom of a basidiomycete.
tail (1) The cytoplasmic portion of a transmembrane receptor teleosemantics A theory of the semantic significance, or meaning,
molecule, with which signal-transducing proteins associate. (2) of symbols that explains meaning in terms of the evolutionary
Domains of the core histones and linker histones involved in history of those symbols.
protein–DNA and protein–protein interactions, and which are teliospore Thick-walled resting spore where karyogamy occurs in
subject to post-translational modification. some basidiomycetes.
talonid The distal part (i.e. towards the back of the mouth) of the teloblastic growth Growth in which new segments arise during
crowns of mandibular/lower premolar and molar teeth. development from a specific budding zone at the rear of the
tandem fusion Direct fusion of telomeric–centromeric and animal.
telomeric–telomeric chromosomal regions (as in Robertsonian telomerase An RNA-containing enzyme complex that extends
rearrangement). chromosome ends (telomeres) by copying its RNA sequence
tapetum Anther tissue that lines the locules containing the repeatedly into chromosomal DNA. This extension enables
developing male gametophytes (pollen grains) and provides DNA replication of the chromosome ends.
materials necessary for their development. telomere A specialized non-coding region of repeated sequence
taphocoenosis Mixed fossil community of fauna and flora that (about 15 000 nucleotides long) found at the ends of eukaryotic
lived in the place of its later embedding together with a fauna and chromosomes. It is replicated in a way that counteracts the
flora transported from other biotopes into the sedimentation tendency of the chromosome otherwise to be shortened at each
area. round of replication.

241
Glossary

telson A posterior projection on the opisthosoma of certain terminator (1) A DNA site which, upon being bound by a
chelicerates. It takes the form of a tail-like spine in horseshoe terminator protein, causes arrest (or pausing) of a replication
crabs, a venomous sting in scorpions and a sensory ‘whip’ in fork when it approaches from one side of the complex, but not the
whipscorpions. other. (2) Signal sequence that halts formation of an RNA
temnospondyls The most abundant and diverse group of archaic transcript and allows for addition of a poly(A) tail.
amphibians. They range from early Carboniferous to early terminus region A restricted and defined segment of the chromo-
Cretaceous and are common in many Triassic fossil assemblages, some in which replication forks meet and fuse.
and were often of large size. The group includes many lizard-like ternary complex (1) A three-component complex, usually con-
forms. sisting of an enzyme protein to which are bound two ligands, such
temperate bacteriophage, temperate phage A bacteriophage as two substrates, or substrate and cofactor or inhibitor. (2) In
that can either cause a lytic infection or persist within its host in a respect of translation, refers to the initiation complex that
prophage state. contains the following three components: eIF2, Met-tRNAi and
temperate virus Virus able to persist within the host cell as a GTP.
provirus integrated within the cell’s DNA. terrestrial Living on the ground.
template A nucleic acid sequence that is copied by an RNA or tertian Associated with fever peaking every 48 hours.
DNA polymerase to produce a new RNA or DNA strand of the Tertiary period The geological period spanning the interval 65
exact complementary sequence to the original sequence. million to about 2 million years ago, characterized by the rise of
temporal cospeciation Strictly contemporaneous cospeciation. mammals.
tension A lengthening of a structure that results from an applied tertiary structure The complete three-dimensional conformation
force (positive linear strain). of a protein.
teratogenic Describes a substance that, when ingested, may cause test A skeletal capsule in some protozoa such as foraminifera and
malformation of the fetus. radiolaria, generally covered by an outer layer of living tissue.
terbinafine A member of the allylamine class of antifungal drugs testa Seed coat.
currently used to treat superficial infections. testicond Retaining the testes in the abdominal cavity rather than
tergite The dorsal cuticular plates of arthropods. A single tergite in a scrotum.
usually surmounts each segment. testosterone A steroid hormone important in the regulation of
terminal complexes The cellulose synthetase particles present on masculine anatomy and sexual behaviour.
the plasma membrane that produce cellulose microfibrils. tetanus High-frequency repetitive action potentials, arising either
terminal deoxynucleotidyltransferase (TdT) A template-inde- from normal neural synaptic activation or from experimental
pendent DNA polymerase that adds non-templated nucleotides stimulation.
at the junctions between gene segments during immunoglobulin tetrad The group of four haploid products of the meiotic divisions
and T-cell receptor gene rearrangement, thus generating addi- or the immediate derivatives thereof. Often applied to the four
tional functional diversity. spores generated by a single meiosis in fungi.
terminal duct lobular unit (TDLU) The peripheral secretory unit tetraether lipid A membrane lipid in which two acyl (phytanyl)
of the breast, composed of a terminal duct, and a lobular chains extend between the inner and outer faces of the bilayer and
arrangement of complex anastomosing tubules embedded in a which are joined to a polar head group on each face by ether
specialized connective tissue stroma. linkage.
terminal oxidase Collective term for cytochrome and quinol tetrahydrobiopterin Essential cosubstrate for phenylalanine
oxidases at the ends of electron transport chains that transfer hydroxylase, tyrosine hydroxylase and tryptophan hydroxylase.
electrons to oxygen, reducing it to form water. It is also referred to as the pterin cosubstrate or, incorrectly, as
terminal rate, terminal velocity The final descent speed of a the pterin cofactor.
particle in a fluid, e.g. a person free-falling in air or a cell in a tetrapods Vertebrate animals with pelvic and shoulder girdles and
liquid. At the terminal rate the fluid resistance equals the net four limbs, including their descendants in which these structures
weight of the particle. have been secondarily lost (as is the case in snakes).
terminal redundancy The presence of identical nucleotide tetravalent Having four binding sites.
sequences at both ends of a linear genome. TH cell Helper T cell. A type of CD4 T lymphocyte that cooperates
terminal repeats Repetitive units of DNA sequence found at the in initiating antibody responses.
ends of linear viral genomes. thalamus A collection of nuclei in the dorsal part of the
terminalization In genetic recombination, the movement of the diencephalon, which relays information to the cerebrum.
chiasmata to the ends of the chromosomes. thallic development In fungi, formation of conidia by septation
termination codon One of three codons (UAA, UAG, UGA) that of a conidiogenous cell, which can enlarge after the conidial
signal the end of a polypeptide chain. initials are delimited.
termination Often refers to the termination of polypeptide chain thalloblastic development In fungi, formation of conidia by
synthesis, triggered by the presence of one of the three septation of a somatic hypha or conidiophores into discrete
termination codons, UAA, UAG or UGA, in the A site. Their conidial initials, which then enlarge.
presence results in the binding to the ribosome of a release factor thallus (plural thalli) The somatic fungal or algal body.
instead of an aminoacyl-tRNA. thanatocoenosis Mixed fossil community of fauna and flora that
terminator protein A protein which, upon binding to a specific lived in the place of its later embedding.
site in the DNA known as a terminator, causes arrest (or pausing) theca A secreted, tightly fitting yet flexible covering around
of a replication fork when it approaches from one side of the some unicellular microorganisms, thus distinguishable from an
complex, but not the other. inflexible, rigid, secreted shell or test. See also lorica.

242
Glossary

thecodont Describes a tooth that sits in a socket in the jaw. thylakoid (1) A simple membrane structure that is the photo-
thecodonts Triassic archosaurs that were the dominant predators synthetic apparatus in cyanobacteria and chloroxybacteria. (2)
throughout the Triassic. Often of large size, they were quad- Vesicles, often stacked, formed from the inner membrane of
rupeds with a sprawling gait. chloroplasts, containing the pigments and enzyme systems
thermal conductance A measure of the propensity of heat to enter essential for photosynthesis.
or leave a body as a function of the thermal gradient between the thymine Pyrimidine base, present in DNA but not in RNA.
body and environment. Thermal conductance is the inverse of thymus An organ located in the chest cavity behind the sternum. It
insulation. is the site of T-cell development.
thermal fluctuations Spontaneous variations in the energy of a thymus-dependent antigen An antigen, e.g. most proteins, that
system due to energy exchange with a surrounding temperature requires an interaction between antigen-specific T cells and B cells
bath. to induce an antibody response.
thermal stability (1) The resistance of DNA or RNA duplexes to thymus-independent antigen Antigen that can induce an anti-
dissociation by increased temperature. (2) The ability of a protein body response without the help of T cells. Such antigens have
to resist denaturation, or unfolding by increased temperature. repetitive epitopes and are able to crosslink the B-cell antigen
thermoduric Heat tolerant. receptor efficiently.
thermodynamic cycle A process that compares the effects of two thyroid-stimulating hormone (TSH) A glycoprotein hormone
single changes separately with the total effect of making both released by the anterior pituitary which stimulates thyroid
changes at once. hormone production and thyroid cell division.
thermodynamics The physics of the relationship between heat thyrotoxicosis The clinical signs and symptoms caused by
and other forms of energy. an excess of the thyroid hormones thyroxine and triiodothyr-
thermophile A microorganism whose optimum temperature for onine.
growth is high, i.e. over 501C. Some thermophilic prokaryotes can Ti plasmid Large tumour-inducing plasmid of Agrobacterium
survive at temperatures approaching 1001C. tumefaciens, responsible for crown gall induction.
thermophilic organism A species that is able to live at high Tic Protein translocation machinery at the inner envelope of
temperatures, i.e. over 501C. chloroplasts.
Thermoplasma A genus of thermophilic archaea that lack cell tiering Development of a vertical ecological stratification in a
walls. forest between the understorey and the canopy.
Thermotoga maritima Hyperthermophilic, rod-shaped marine
till Sediment produced by glacial action, characterized by coarse
bacterium with a coat-like cell envelope (‘toga’).
stratification, angular fragments, and a wide range of particle
theropod dinosaur Meat-eating dinosaurs that walked on their
sizes from silt to boulders.
hind feet. Examples include Allosaurus, Velociraptor and
tillite A sedimentary rock produced from glacial till.
Tyrannosaurus.
TILs See tumour-infiltrating lymphocytes.
Theropoda See theropod dinosaurs.
tinamous A taxon of fowl-like birds found in semi-arid environ-
thioester Compound (R–CO–S–R0 ) in which a carboxylic acid
ments which have many characters of the Palaeognathae and
(R–COOH) is condensed with a thiol (R0 –SH).
therefore systematically are generally placed in or very near to
threes A set of three ciliates united in conjugation, in contrast to
that taxon.
the normal pair formation during conjugation.
threshold (1) The level that has to be reached by a substance or tip growth A mechanism of growth of plant cells in which new wall
stimulus before it will trigger a subsequent event. (2) In excitable material is added only to one end of certain elongating cells, such
cells, the membrane potential at which sodium channels open as pollen tubes and root hairs.
rapidly, depolarizing the cell and causing an action potential. tissue A coherent assembly of cells serving a particular function in
Experimentally, the threshold is determined as the membrane an organism.
voltage reached by a stimulus that generates an action potential tissue cyst Enlarged host cell containing numerous parasites with
during 50% of trials. reduced activity. It may be strengthened by primary and/or
thrombin A central protein in haemostasis, generated from its secondary cyst walls.
inactive precursor prothrombin. Thrombin converts fibrinogen tissue factor A single-chain glycoprotein complexed to phospho-
to fibrin, but can also cause cellular activation of platelets, lipid that is the main physiological activator of blood coagula-
monocytes and endothelium. tion. It is expressed on cells within the vessel wall, but not
thrombocytopenia A decrease in the number of circulating normally on cells in direct contact with the blood.
platelets in the blood. Tithonian The top stage of the Upper Jurassic.
thrombogenic A substrate that induces platelet aggregation and titre (1) A measurement of the dilution at which an antiserum can
activation of the plasma coagulation factors. be detected in an assay. (2) Number of infectious virus particles
thrombomodulin A protein expressed on the endothelium of per given volume of a virus stock solution.
blood vessels, which can form a 1:1 stoichiometric complex with TLX See trophoblast leucocyte cross-reactive antigen.
thrombin and thus alter its substrate affinity. The complex of Toc Protein translocation machinery at the outer envelope of
thrombin–thrombomodulin activates protein C. chloroplasts.
thrombophlebitis Inflammation of a vein leading to a blood clot. tolerance Failure to respond to an antigen that is, in other
thrombosis The formation of an intravascular blood clot (throm- conditions, capable of inducing an immune response. Self
bus), consisting of fibrin, aggregated platelets and blood cells, in tolerance is an essential feature of the immune system and is
response to an imbalance between thrombogenic factors and established by the deletion/inactivation of self-reactive clones
protective, anticoagulant mechanisms. during lymphocyte development.

243
Glossary

tolerogenic Able to induce specific immunological ‘tolerance’ trachea (plural trachaeae) (1) The windpipe in vertebrates. (2) In
(non-reactivity). The test for the tolerant state is to observe zero arthropods, a thin-walled cuticular tube that opens through small
response following challenge with antigen in a form that would be holes in the body surface and extends throughout the body,
expected to stimulate an immune reaction. providing tissues with atmospheric oxygen and a pathway for
Toll-like receptors A family of mammalian transmembrane removing carbon dioxide.
receptors related to the Toll protein of Drosophila. They are tracheid A water-conducting xylem tissue in plants, composed of
involved in the recognition of pathogens and microbial products tapering tracheary elements that do not have open ends.
and activate antimicrobial effector pathways in phagocytes. trajectory The sequence of points (in the phase space) visited by a
tomography Any technique that allows ‘slices’ of the body to be system as time moves forwards.
imaged, with the partial or complete elimination of effects from trans-acting Acting on or affecting another molecule.
all other regions of the body. Image data are acquired from a trans-acting determinant A gene product that influences the
number (often large) of different perspectives round an object, expression of another gene.
whereby a cross-sectional view of it can be produced. trans-acting factor A macromolecule or complex of macromole-
tonoplast Membrane surrounding the vacuole in plant and fungal cules (usually protein and/or RNA) that interacts with RNA (or
cells. DNA).
tool kit Term used by archaeologists to refer to the various types of transamination The transfer of an amino group from an amino
tools found in ancient tool caches, or in circumstances where acid to an a-ketoacid to produce a new keto acid and a new amino
natural agencies have brought stone tools together. acid.
topographic projections The situation when axons of neighbour- transcript An RNA molecule produced by transcription.
ing neurons innervate neighbouring areas in their target field. transcriptase DNA-dependent RNA polymerase. An RNA poly-
topography A detailed surface map of a region. merase that catalyses the formation of RNA on a DNA template.
topoisomerase Class of enzymes involved in controlling the coiled transcription The synthesis of RNA molecules using DNA as the
or relaxed state of DNA by catalysing transient breaks. Type I template to determine the sequence of bases in the RNA product.
topoisomerases catalyse single-strand breaks and type II double- The synthesis of RNA is catalysed by the enzyme RNA
strand breaks. polymerase.
topology In regard to phylogenetics, the connectivity relationship transcription factor A regulatory protein required to initiate,
of nodes in a phylogenetic tree. Trees with the same topology may upregulate or repress transcription. The term originally referred
have different branch lengths and may be drawn differently, but especially to those factors involved in the precise binding of RNA
represent the same branching pattern. polymerases to promoters on the DNA and the initiation of
transcription, but is now widely used for any gene regulatory
top-up transfusion Small amounts of blood given to an infant to
protein.
increase the haemoglobin concentration. The neonate can
transcription-coupled nucleotide-excision repair A DNA re-
tolerate only 10 ml kg1 of concentrated red cells. If it is necessary
pair process by which nucleotide-excision repair occurs prefer-
to give more blood, an exchange transfusion has to be performed.
entially on the transcribed strand on a transcriptionally active
toroid In the shape of a torus, like a doughnut with a hole in the
gene, presumably by the coupling of repair and transcription.
middle.
transcriptome The full complement of RNA species transcribed
torpor A regulated state of inactivity where the body temperature
by a cell.
is regulated somewhere in the region between 01C and
transcytosis A transport process by which molecules cross a
euthermy.
cellular barrier in a membrane-bound, or pinocytic, vesicle. The
torque The turning or twisting force, the product of the magnitude
process involves pinocytic uptake, or endocytosis, of the molecule
of a force, F, and the perpendicular distance between the line of
at one surface of the cellular barrier, and pinocytic release, or
action of the force and the centre of rotation, d.
exocytosis, at the other.
torsion angle The angle between two groups on either side of a transdifferentiation Differentiation of one cell type into another.
freely rotating chemical bond. transduce To infect cells with a virus.
torsion A counterclockwise 1801 rotation of the visceral part of transducer (1) A device for converting one form of energy into
the body in gastropod development and phylogeny. It is the another. Biologically, a device for converting the energy of a
diagnostic character of the class Gastropoda. stimulus applied to a tissue into a series of action potentials in an
torus semicircularis Also known as the inferior colliculus in afferent nerve. (2) A device for converting a signal from one form
mammals. It is part of the tectum (or roof) of the midbrain that into another, e.g. a cell-surface receptor, which transmits the
receives auditory input and, where present, lateral line input. signal that extracellular ligand has bound to an intracellular
toxicity The intrinsic property of certain substances to be harmful signalling pathway. (3) The region of a sensory protein
to the normal structure or physiology of cells, tissues or organs. responsible for signal generation.
Toxoplasma A protozoan parasite causing a disease of animals and transduction (1) Signal transduction. The process of translating a
humans, transmitted from cats (the primary host). stimulus into an intracellular response. (2) Virus-mediated
toxoplasmosis Disease caused by infection with the protozoan transfer of a gene into a cell.
parasite Toxoplasma gondii. transencapsidation The packaging of one type of virus in the
TPs See transition polypeptides. other’s protein coat, when two different viruses infect a cell.
trabecula A solid strand of connective tissue branching from the trans-encoded antisense RNA An RNA encoded by a gene that is
splenic capsule into the interior of the spleen. not linked to its target gene. Target and trans-encoded antisense
trace fossil Evidence of activity of an ancient animal or plant, such transcripts are not completely complementary and form im-
as a burrow, track or faecal pellet. perfect RNA–RNA duplexes.

244
Glossary

transesterification The reaction where one ester moiety is transition-state analogue inhibitor A stable molecule that binds
replaced by another. to an enzyme in a manner that is structurally analogous to the
transfected cell A cell into which a bacterial plasmid that contains transition state.
a foreign virus or genetic material has been inserted. translation The process whereby the nucleotide sequence of a
transfer RNA (tRNA) A family of small RNA molecules that act as messenger RNA is read out and used to make a polypeptide
‘adaptors’ in the process of translating the sequence of a chain. It takes place on the ribosomes. The process is called
messenger RNA into protein. Each tRNA molecule carries an translation because the alphabet of nucleic acids (AT/UGC) is
amino acid that matches its anticodon (which will match to the converted into sequences of amino acids.
appropriate codon in mRNA). tRNAs are small (75–100 translational control Regulation of protein synthesis at the
nucleotide) elbow-shaped RNA molecules that carry a three- translational stage.
base sequence (‘anticodon’) on the long arm and an amino acid on translational operator Sequence in mRNA, generally encom-
the short arm. passing the translation initiation region, to which translational
transferase An enzyme that breaks a bond in a donor substrate repressor proteins bind.
and transfers one group from it to an acceptor substrate. The translational polarity The fact that translation of a message on a
remainder of the donor substrate is the ‘leaving group’. polycistronic messenger RNA depends on the translation of a 50
transferrin A transport protein in animals that binds, transports preceding message.
and delivers free iron to cells. translational repressor Protein which binds to an mRNA, usually
transform In image processing, to take an image from one space near the translation start, blocking access of ribosomes and
(domain) to another. The most commonly used transform in inhibiting protein synthesis.
image processing is the Fourier transform which takes an image
translocase A protein or proteins that promotes movement
from coordinate space to frequency space.
(translocation) of macromolecular machinery (e.g. ribosome or
transformation (1) The process by which bacteria and yeast cells
spliceosome) along an RNA.
take up exogenous DNA. (2) A multistep process involving
translocation (1) The step in the elongation cycle of protein
mutation of cellular genes or viral infection by which cells acquire
synthesis during which A- and P-site tRNAs move to P and E
altered growth properties, usually including the ability to divide
sites, respectively, and the messenger RNA is advanced by one
indefinitely in culture. The pheotype resembles that of cells from
codon. (2) A change in location, as in the movement of a molecule
cancerous tissue.
from one compartment of the cell to another. The term often
transgene A piece of foreign DNA that has become incorporated
refers specifically to the movement of proteins across a
into the genome of a cell.
membrane. (3) A chromosome rearrangement in which there is
transgenesis The intentional introduction of foreign genetic
exchange of parts between nonhomologous chromosomes.
material into the genome of plants or animals.
transmembrane Extending completely across a lipid bilayer, with
transgenic Carrying a foreign gene. Describes an organism in
exposure on both sides of the bilayer to the adjacent aqueous
which the deliberate insertion of a foreign gene into the germ cells
compartments.
or into the very early embryo has given rise to a mature organism
with the foreign transgene incorporated into the genome of every transmembrane helical domain (TM) a-helical secondary struc-
cell. ture in those parts of the protein chain that cross the membrane in
transgenic animals Animals carrying and expressing a foreign integral membrane proteins.
gene. transmembrane receptor Cell-surface protein whose
transgenic mice Genetically engineered mice that contain a extracellular domain recognizes and binds an extracellular
foreign gene integrated into the germline genome. signal molecule, and which transduces this information into
transgenic plant A genetically engineered plant that contains and the interior of the cell by association of the cytoplasmic
expresses a modified gene or a gene from another species. portion of the receptor with intracellular signalling
transglycosylation reaction The insertion of a base by cleavage of machinery.
the N–C glycosidic bond between the nucleobase and the C1 of transmembrane synthases Enzymes located in the plant cell
ribose. membrane that extend from the cytoplasmic side of the
transglycosylation Cleavage of a glycosidic bond followed by membrane to the external, environmental, side and that are
attachment of the newly formed (potentially) reducing terminus active in synthesizing cell wall components.
to some other molecule (e.g. another sugar). transmissible spongiform encephalopathies (TSEs) Diseases
transgression The drowning of the land by a relative rise in sea caused by defective brain proteins, called prions, that lead
level. to brain degeneration. In humans, they include Creutzfeldt–
transition See transition mutation. Jakob disease (CJD), Gerstmann–Sträussler–Scheinker
transition mutation A mutation in which a purine (adenine, syndrome (GSS) and fatal familial insomnia (FFI). In animals
guanine) replaces another purine, or a pyrimidine (cytosine, they include bovine spongiform encephalopathy (BSE) and
thymine) replaces another pyrimidine. scrapie.
transition polypeptides (TPs) Set of small proteins with mass transmitter Chemical substance liberated from presynaptic axon
33 kDa that are synthesized when a plant is initially exposed to terminals when they are invaded by propagating action
an anaerobic environment. For the first 90 minutes, these are potentials. It acts on receptors on the postsynaptic cell to
the only proteins that appear to be synthesized. generate postsynaptic potentials, thus passing the signal from one
transition state A short-lived, high-energy intermediate molecular cell to the other.
state that must be attained in order to complete the transforma- transovarial Transfer of infection from the adult female tick to the
tion of reactants to products in any chemical reaction. next generation via the eggs.

245
Glossary

transparent testa Arabidopsis phenotypes recognized by the trigeminovascular reflex Link between nerve cells and vascular
reduction or absence of pigments in the seed coat (testa), due to mechanisms such that stimulation of nerve cells in the brainstem
the absence of one or more loci required for flavonoid via the parasympathetic part of the facial nerve increases blood
biosynthesis, thus revealing the yellow colour of the underlying flow to the head.
tissues. trilete mark Triradiate feature on a plant spore through which
transplantation (1) The surgical implantation of tissue or an organ germination occurs.
taken from one individual (the donor) into another individual trimeric Describes a macromolecule that is composed of three
(the recipient). (2) Insertion of a few cells from a donor embryo subunits.
into a host of the same age (homochronic) or of a different age triple gene block Three contiguous virus genes located at the 30 -
(heterochronic). terminus of the genomic nucleic acids of potexviruses and
transport Often refers to membrane transport, the transport of carlaviruses.
substances, typically small molecules and ions, across a trisomy A condition of having three copies of a given chromosome
membrane, instead of the normal two.
transporters Transmembrane proteins that transport solutes trivalent The association of three chromosomes during meiosis I
or ions across the membrane, using a variety of energy metaphase, typically observed in Robertsonian translocation
sources. heterozygotes.
transporter multimers Functional transporter units consisting tRNA See transfer RNA.
of more than one copy of the transporter protein. This is not tRNAMetf Initiator tRNA not carrying its amino acid.
established or ruled out for plasma membrane transporters. tRNAMetf Initiator tRNA in prokaryotes. The methionine amino-
transposable element A DNA sequence that is able to move acylated to this tRNA is formylated.
itself, or a copy of itself, to a new location in the genome (the tRNAMetf/m tRNA from animal mitochondria, which occurs as
process of transposition). Such movements often cause muta- fMet-tRNAMet Met
f/m and Met-tRNAf/m . The formylated form is
tions. active in initiation of translation and the non-formylated form
transposase The catalytic protein that carries out the DNA in the elongation of protein synthesis.
cleavage and strand-transfer reactions of transposition. tRNAMeti Initiator tRNA in the eukaryotic cytoplasm.
transposition A process whereby certain DNA elements (insertion The methionine aminoacylated to this tRNA is not
elements and transposons) move from one DNA location to formylated.
another, using enzymes (transposases) usually encoded by the tRNAMetm The methionine-carrying tRNA that acts during elonga-
element. tion of the polypeptide chain.
transposon See transposable element. trophectoderm Simple squamous epithelium that covers the
trans-splicing Formation of a mature messenger RNA by splicing blastocyst. Its derivatives are giant cells, the ectoplacental cone
of exons from different primary RNA transcripts. It is contrasted and extraembryonic ectoderm, all of which contribute to the
with the more usual cis-splicing, in which exons from a single chorionic disc of the placenta.
transcript are spliced together. trophic factors Proteins that support the survival and growth of
TransTerm A database of translational signals including initiation cells.
and termination contexts from more than 250 organisms and trophic level The place of an organism in a food chain.
currently containing nearly 100 000 sequences. trophonemata Vascularized villous extensions of uterine mucosa
transversion See transversion mutation. in stingrays used for nutrient delivery and respiration.
transversion mutation A mutation in which a purine (adenine, trophozoite The feeding, motile stage of a sporozoan protozoan.
guanine) replaces a pyrimidine (cytosine, thymine) or vice versa. trypanosomiasis Disease caused by a protozoan parasite of genus
treadmilling A description of the condition in which monomers Trypanosoma.
are being added to one end of a filament at the same time as they TSEs See transmissible spongiform encephalopathies.
are depolymerizing from the other end. tsetse fly Dipteran fly of genus Glossina restricted to sub-Saharan
triacylglycerol Ester of glycerol with three fatty acyl side chains. Africa.
triangulation number A description of a triangular face of an TSH receptor Receptor for thyroid-stimulating hormone. It is a
icosahedron indicating the number of triangles into which each member of the G-protein-coupled receptor family.
face is divided when forming an icosadeltahedron. TSH See thyroid-stimulating hormone.
Triassic period First major division of the Mesozoic era. tsunami A great sea wave produced by submarine earth movement
triazole An azole antifungal agent containing an azole ring with or volcanic eruption.
three nitrogen atoms. Examples are fluconazole and itraconazole. TTV1 Virus that infects the archaeon Thermoproteus tenax.
trichobothrium (plural trichobothria) Long, thin, hair-like tubulin The protein monomer that polymerizes to make micro-
structure with a jointed attachment to the cuticle in arthropods. tubules.
Used in many terrestrial arthropods for sensing slight displace- tuff Consolidated ash derived from a volcanic eruption.
ments or vibrations of the air. Tullgren funnel A funnel used for extracting soil fauna. The funnel
trichocyst An organelle in the cell cortex which is capable of holds a soil sample on a wire mesh and this is heated from above
undergoing an explosive change in shape to extrude a (generally with an incandescent bulb to drive animals into a collecting vessel
thread-like) projection in response to stimulation. below.
trichogyne A slender hypha originating from the gametic cell of a tumour suppressor A recessive gene normally involved in
protoperithecium that will fuse with a cell of the opposite mating preventing cell proliferation, which, when eliminated at both
type during fertilization. alleles, allows uncontrolled proliferation.

246
Glossary

tumour-infiltrating lymphocytes (TILs) T cells isolated from a ultrasensitive response Also called a sigmoidal response. A
tumour which, after treatment with interleukin-2 to stimulate cell response described by an S-shaped stimulus–response curve.
proliferation, can mediate tumour regression in some cancers Ultrasensitive responses are more switch-like than Michaelian
when reintroduced. responses.
tunnelling The contribution made to chemical reaction rates by ultrasonography The use of high-frequency sound (usually 1–
virtue of the wave character of matter. 10 MHz range) to generate images of the human body.
turbinates Scroll-shaped bones in the nasal passages of birds and ultrastructure The structure of a cell that is only visible under a
mammals. high-power optical microscope or an electron microscope.
turgor A positive internal hydrostatic pressure that develops in uncoating The stage of viral replication at which structural
plant cells as a result of the osmotic intake of water and the proteins are lost and the virus genome is exposed to the
presence of the rigid plant cell wall. It is an important driving replication machinery.
force for cell growth. underdominance The lower fitness of heterozygotes relative to
turgor pressure The pressure exerted on the plasma membrane in either homozygote.
a plant cell due to the osmotic differences between the cytoplasm underreplication The failure to completely replicate the DNA of a
and the outside environment. chromosome.
undulating membrane In certain protists, a fold of the cell
twitching motility A form of surface translocation in bacteria that
membrane united with the flagellar membrane, resulting in
is attributable to type IV fimbriae.
attachment of the flagellar axoneme to the cell surface. In the
two-component system See two-component regulatory system.
trichomonads it results in the formation of the recurrent
two-hybrid system A system for examining protein–protein flagellum.
interactions in vivo. It involves specialized genetic constructs unequal exchange Recombination between tandemly repeated
transformed into yeast cells, and can be used to test interactions sequences that are aligned out of register.
between yeast proteins or to study other proteins expressed in unfolded The state of a protein molecule that exists in a large
yeast. number (ensemble) of random conformations that are inter-
type A specimen that is irrevocably tied to that species name. converting. Unfolding is often induced by chemical denaturants
type I hair cells, type II hair cells Hair cells in the vestibular or increased temperature.
system, which have or lack a calyx, respectively. unguligrade A foot posture in which the animal stands on the last
typology See essentialism. phalanx of the digit – on tip toes – with the sole or palm of the foot
tyrosinase See phenooxidase. and the digits off the ground.
tyrosine kinase Any enzyme that phosphorylates certain proteins uniform binding Binding that arises from an enzymatic interac-
on one or more tyrosine residues. tion that comparably stabilizes all internal states.
tyrosine phosphorylation A mechanism for activating or Uniform Resource Locator See URL.
inactivating proteins by phosphorylation of specific tyrosine unilocular Describes a sporangium formed of a single cell, which
residues. undergoes meiotic division commonly followed by mitosis after
tyrosyl-tRNA synthetase Enzyme that attaches tyrosine to its which the cytoplasm divides to produce four or more spores
cognate tRNA, in a reaction that links the carboxyl group of (usually motile).
tyrosine in an ester linkage to the phosphate of AMP. Polymers of uniparental disomy The situation where both copies of a
glycerol and ribitol joined by phosphate groups, with amino acids particular chromosome are inherited from one parent and none
or sugars attached. Major components of the Gram-type positive from the other.
cell wall, these polymers are found in great variety and are highly uniporter A transport protein (carrier) that facilitates passive
antigenic. diffusion of a solute across a membrane.
unipotent stem cell A stem cell that gives rise to a single type of
differentiated cell.
unit cell In crystallography, the representative parallelepiped
U1 snRNA A highly conserved small nuclear RNA involved in defined within the crystal, which may be tessellated with itself
splicing. The U1 snRNP functions in 50 splice site recognition in three dimensions to build up the complete crystal.
during splicing. unmyelinated fibres Nerve cell axons that are wrapped by glial
U2 auxiliary factor (U2AF) A dimeric protein containing (or Schwann) cells but without the formation of a myelin sheath.
RNA-binding and RS domains that associates with unpaired electron An electron that is the sole occupant of an
pre-mRNA near the 30 splice site through direct binding of atomic or molecular orbital.
the pyrimidine-rich sequences and through protein–protein unsaturated fatty acid Fatty acid that contains one or more
contacts. double bonds in the hydrocarbon tail that can accept hydrogen or
U2 snRNA A highly conserved small nuclear RNA involved in other substituents.
splicing. The U2 snRNP functions in branch site recognition untranslated region (UTR) The untranslated region of an
during splicing. U2 remains associated with the pre-mRNA mRNA, located either 50 (50 -UTR) or 30 (30 -UTR) to the coding
throughout splicing and may form part of the catalytic core of the region.
spliceosome. upstream Toward the 50 end, a term used to indicate direction
ultracentrifuge A high-speed centrifuge capable of reaching up to along a DNA or RNA strand. Transcription proceeds from
200 000 g. upstream to downstream on DNA (50 to 30 ).
ultramafic Rocks that are very rich in magnesium and iron, such as upwelling An area in the ocean where water is moving vertically
peridotites. towards the surface.

247
Glossary

uracil A commonly occurring pyrimidine base, found in RNA. variable regions The N-terminal domains of antibody heavy and
urbilaterian The primitive bilaterian from which the protostome light chains. They vary in amino-acid sequence between different
and deuterostome lineages diverged. clones and form the antigen-binding site.
urea CON2H4, a non-toxic product of nitrogen waste formed by varicella-zoster An acute infectious viral disease, known com-
the urea cycle in the liver. monly as chickenpox. The same virus also causes shingles.
urea cycle A metabolic pathway in the liver in which amino groups vas deferens A muscular duct that carries sperm from the
are donated by ammonia and aspartic acid and combine with epididymis to the ejaculatory duct.
carbon to form urea. vascular addressins Tissue- or organ-specific endothelial cell-
ureotelic Describes organisms that excrete nitrogen principally in surface molecules that are ligands for homing receptors on
the form of urea. lymphocytes, and which direct the migration of lymphocytes into
uricotelic Describes organisms that excrete nitrogen principally in the appropriate target tissue.
the form of uric acid. vascular bundle In plants, a discrete bundle of xylem and phloem
URL Uniform Resource Locator, an information source described providing a conduit for nutrients and water.
by the communications protocols (http), the Internet address of vascular cambium A band of meristem around the central
the host computer and the file location of the specific informa- vascular tissue that produces secondary vascular tissues in woody
tion. plants, which is is responsible for the growth in girth of trunks and
uropod The most terminal posterior appendage on the crustacean branches.
abdomen. vasculitis An inflammatory disorder affecting blood vessels and
urticaria Hives, an allergic skin condition characterized by dermal resulting in destruction of vessel walls and reduction of blood
oedema (wheal) and surrounding erythema (redness) as a result of flow to tissues.
the release of histamine and other mediators from mast cells in the vasoactive Refers to an agent that exerts an effect on the diameter
skin. (calibre) of blood vessels. Vasoconstrictive agents decrease
calibre and slow blood flow, and vasodilative agents increase
utilization See extraction.
calibre and flow.
UTR See untranslated region.
vasoconstriction A reduction in the cross-sectional area of a blood
U-turn motif A structural motif first found in tRNA anticodon
vessel.
loops. The loop structure is highly constrained by unusual
vasoconstrictor An agent that causes narrowing of the blood
hydrogen-bond interactions within the loop.
vessels.
uveitis Inflammation of the middle epithelial layer (uvea) of the iris
vasodilation, vasodilatation An increase in the cross-sectional
of the eye.
area of a blood vessel.
vasodilator-stimulated phosphoprotein (VASP) An adaptor
protein that binds to ActA, zyxin and viculin through one
Vmax The maximum steady-state velocity achievable by an enzyme proline-rich domain and to profilin through a second proline-rich
at infinite substrate concentration. It is also equivalent to the domain.
value of kcat multiplied by the enzyme concentration. vasomotor responses Constriction (vasoconstriction) or relaxa-
vac genes Genes associated with formation of gas vesicles in tion (vasodilation) of smooth musculature of blood vessels, often
bacteria. with reference to peripheral arterioles.
vac operon An operon of gas vesicle genes. VASP See vasodilator-stimulated phosphoprotein.
vaccination Administration of a pathogen-derived antigen or the Vb Any of the V gene segments at the T-cell receptor b-chain locus.
whole microorganism (in killed or non-pathogenic form) in order The final T-cell receptor will contain one Vb sequence.
to produce an antigen-specific immune response that will protect VDJ recombination The somatic recombination of germline-
against the disease. encoded gene segments (V and J in the case of immunoglobulin
vaccine Originally, the cowpox material used by Edward Jenner to light chains and T-cell receptor a chains, and V, D and J in heavy
immunize against smallpox. The term is now used for any chain and b chain) to generate a sequence encoding a functional
preparation used for preventive immunization against a specific variable region. It occurs only in developing T cells and B cells and
disease-causing microorganism. is a key event in their development.
vadose zone The unsaturated zone above the water table. This vector (1) A living agent that can act as a vehicle to transmit a
zone may be considerably deeper than the soil and include disease from one host to another. (2) A piece of DNA that is
underlying materials unsaturated with water. capable of propagating itself inside cells and is used to carry DNA
valve One half of the two-part shell of some molluscs (e.g. clams sequences of interest in cloning and gene expression experiments.
and cockles) and brachiopods. A vector that can replicate in different organisms is called a
van der Waals forces, van der Waals interaction A relatively shuttle vector. A vector that carries the foreign DNA in a form in
weak interaction between any pair of atoms that are in close which it can be expressed is called an expression vector.
proximity to each other. They show a weak bonding interaction vegetative cell The growing cell that, in the case of a typical
due to their fluctuating electrical charges. The strength of the prokaryote, divides by binary fission at a rate corresponding with
interaction falls off rapidly as the distance between the atoms is the level of nutrients.
increased. velocity In enzyme kinetics, the rate of an enzyme-catalysed
varanids A group of mostly large, predatory lizards that include reaction, defined as the change in substrate or product
the living monitors. They are characterized by light flexible skulls, concentration per unit time.
long necks and retractable, forked tongues. velvet worms See Onychophora.

248
Glossary

venomous animal Animal that produces and excretes toxic virulence A measure of the harm that a pathogen can inflict on its
compounds. host.
ventral Front side or underside of a structure or organism virulent phage A phage that produces a lytic infection and cannot
(opposite to dorsal). Towards the belly of a vertebrate. enter a prophage state.
vermiform Worm-shaped. virus attachment protein The protein on the surface of a virus
vertebrates Animals, including fishes, amphibians, reptiles, birds particle responsible for binding the receptor.
and mammals, that possess a dorsal nerve cord supported by a virus-like particles (VLPs) Particles that self-assemble following
cartilaginous or bony vertebrae (the backbone). Phylum Chor- overexpression of the virus capsid protein in a baculovirus
data, subphylum Vertebrata. expression system. The VLPs are antigenically authentic and
vertical transmission (1) The transfer of symbiotic microorgan- morphologically resemble virus particles but contain no nucleic
isms from parent to offspring, often by direct insertion into (or on acid.
to) gametes or other reproductive propagules. (2) Transmission viscera The internal organs, such as heart, liver, intestines.
of disease from mother to offspring via infection in the womb. visceral endoderm Extraembryonic cell lineage which gives rise to
vesicles (1) Small membrane-bound structures in eukaryotic cells the yolk sac endoderm.
that carry material from one compartment to another and to the visceral Pertaining to the internal organs.
plasma membrane for secretion. (2) Internal particles produced visceromegaly See organomegaly.
by bacteria to perform a distinct function. For example, gas
vital rates Birth and death rates, usually age- or stage-specific.
vesicles adjust cell buoyancy, vesicles containing bacterio-
vital stain A dye that is taken up differentially by live and dead cells
chlorophyll are necessary for photosynthesis.
and can be used, for example, to visualize viral plaques.
vesicular systems In muscle fibres, the internal membrane-
vitamin C deficiency A state of malnutrition due to a lack of
bounded compartment from which calcium is released to raise
ascorbic acid (vitamin C) in daily foods.
internal calcium levels to activate the myofilament system.
vesiculoviruses Viruses classified in the genus Vesiculovirus of the vitreous ice Also called amorphous ice. Water that has been frozen
family Rhabdoviridae. in such a way that no crystals have been created. Protein
imbedded in vitreous ice is thought to keep its original structure.
vessel A water-conducting xylem tissue composed of a line of
tracheary elements with open ends. vitrification Cooling a sample rapidly enough to prevent crystal
formation. Vitrification of pure water requires an extremely high
VF See ventricular fibrillation.
cooling speed.
Vg1 A member of the transforming growth factor \beta super-
family of secreted signalling proteins. It is expressed in Xenopus vivipary The production of live young directly from the mother’s
embryos at the vegetal pole. body, either by retention of eggs until hatching or by the
intermediary of a placenta.
viability (1) An individual’s ability to survive, measured as the
probability of surviving from birth to adulthood. (2) Of seeds, the VLP See virus-like particles.
percentage in a sample that germinate to give normal healthy vnf Gene designation of the vanadium nitrogenase system.
seedlings. VNTR See variable number of tandem repeats.
vicariance The separation of a formerly continuous, ancestral volcanic arc A chain of volcanic mountains formed above the
population into two or more new populations as a result of subduction zone by magma rising up from the melting of the
geological or climatic changes. downgoing plate.
vicariance biogeography Approach to biogeography that as- voltage-dependent calcium channel, voltage-gated calcium
sumes that the modern distribution of a species is caused by the channel An ion channel, situated in the plasma membrane, that
splitting (vicariance) of a former more extensive range rather than is selectively permeable to calcium ions and opens in response to
by dispersal events. membrane depolarization.
vigour In plants, a measure of the relative rapidity and uniformity voltammetry Technique by which substances can be measured in
of germination, which results in stronger seedlings and better crop fluid on the basis of oxidation–reduction reactions generated in
establishment. probes engineered for these electrochemical reactions.
viraemia The presence of virus in blood. volutin Metachromatically staining polyphosphate storage gran-
viral oncogene A gene carried by a retrovirus or DNA virus whose ule found in some microorganisms.
product causes cellular transformation in vitro, and/or contri- vomeronasal organ A sensory structure in the nose or roof of the
butes to tumour induction in vivo. mouth of many vertebrates. It responds to specific volatile
viral quasispecies A dynamic distribution of closely related, compounds termed pheromones, which animals use to commu-
replicating mutant and recombinant genomes subjected to a nicate with each other. In snakes it comprises a pair of blind-
process of genetic variation, competition and selection. ended chambers in the roof of the mouth. They are lined with
viral vectors Replication-deficient viral particles that are used as sensory cells and process the molecules picked up by the tongue
vehicles to introduce genetic material into target cells. during tongue-flicking.
Virchow–Robin spaces Extensions of the subarachnoid space voxel The basic volume-element in the MRI/MRSI image being
along blood vessels which penetrate the brain parenchyma. acquired. It is the three-dimensional analogue of the ‘pixel’.
virion A complete virus particle containing the genetic material, VPg A small 15-kDa protein covalently attached to the 50 -end of the
capsid, and envelope (where applicable). It can exist outside living genomic and subgenomic RNAs of caliciviruses.
cells and is capable of infecting cells. vulva Opening in the ventral epidermis of the adult hermaphrodite
viroplasm Inclusion body in a cell where viral replication and nematode, connecting the gonads to the outside. It allows mating
assembly is taking place. and egg-laying.

249
Glossary

wall loosening Proposed interruption of biochemical crosslinks wobble, wobble base pair A nonstandard base-pair interaction
between the load-bearing components of the plant cell wall that between the third nucleotide of a codon (the wobble base) and the
allows wall expansion. first nucleotide of the anticodon. In this way a single tRNA is able
Wallerian degeneration The degenerative changes in the distal to decode multiple codons.
stump of a nerve that occur following axonal damage. Named Wolffian duct The mesonephros duct which subsequently gives
after Waller, who discovered this phenomenon in 1850. rise to the ureteric bud that interacts with metanephric
warm-reactive autoantibody An antibody to red blood cells that mesenchyme to produce metanephros.
binds to the cells most efficiently at 371C. It is principally IgG. work The product of force and distance (measured in joules, J, the
Watson–Crick base pairing, Watson–Crick pairing rules energy needed to move 1 newton (N) a distance of 1 m).
Hydrogen bonding in nucleic acids between A (adenine) and T woven bone Bone tissue that has coarse collagen fibre bundles
(thymine) or U (uracil), and between G (guanine) and C interwoven in varying directions.
(cytosine). Wright–Fisher model A model of reproduction which assumes
wax ester Ester of a long-chain fatty alcohol with a long-chain that the alleles present at a given generation are a random sample
fatty acid. from the gene pool of the previous generation.
WD-40 A repetitive tryptophan–aspartic acid motif in Gb sub-
units.
Web The World Wide Web (WWW), a set of communication xenoma A hypertrophic growth of host cells of microsporidial
protocols with hypertext capability developed at the High- parasites, forming giant cells filled with microsporidial spores
Energy Physics Laboratory, CERN, in Geneva, Switzerland. that are frequently encountered as tumours on fish and other
Weil disease Severe form of leptospirosis with jaundice and renal hosts.
failure. X-ray film A thin piece of plastic covered by silver halide crystals
wet deposition Deposition of (pollutant) material removed from capable of undergoing oxidation by high-energy species and thus
the atmosphere by precipitation. revealing a latent image.
whole-arm fusion (centric fusion, Robertsonian transloca-
tion) Fusion of the long arms of two chromosomes following
breakage and loss of the small arms of two acrocentric YAC Yeast artificial chromosome, containing yeast telomeres,
chromosomes. origin of replication and centromere. There is a multiple cloning
Wiebel–Palade bodies Storage granules present in endothelial site and positive selection for the cloning vector.
cells. They are a component of the secretory pathway of yolk sac Extraembryonic tissues (ectoderm, mesoderm and endo-
endothelial cells. Factor VIII, von Willebrand factor, P-selectin, derm) that surround and digest yolk stores. In some sharks, it
interleukin-8, and endothelin are examples of molecules stored in forms the yolk sac placenta.
these organelles in endothelial cells.
wilderness A large area that remains essentially unmanaged and
unmodified by human beings. zalcitabine Nucleoside analogue inhibitor of human immuno-
wild-type Describes the organism as first isolated from nature, deficiency virus (HIV) reverse transcriptase. Used as an anti-HIV
prior to introduction of additional mutations. In some genetic drug.
studies a natural variant is arbitrarily chosen as the ‘wild-type’ zanamivir A neuraminidase inhibitor that acts as an anti-influenza
point of reference. agent.
window The parameters of grey scale chosen for image viewing. zidovudine Nucleoside analogue inhibitor of human immuno-
The window level is centred on the attenuation of the tissue in deficiency virus (HIV) reverse transcriptase. Used as an anti-HIV
question. The window width refers to the range of other tissues drug.
demonstrated. zonal bone Periosteal bone with ‘growth rings’.
wing-finger Highly enlarged fourth digit of the pterosaur hand Z-test A statistical test of significance that measures how
that supported the outer part the wing. far an observation is from the mean in units of standard
Wiskott–Aldrich syndrome Genetic disease characterized by error.
eczema, thrombocytopenia and recurrent infections. zygote The immediate product of fusion of sperm and egg
witches’ broom Symptoms of abnormal, massed, brush-like at fertilization. It comprises a single cell that contains
development of shoots or roots, mainly on woody plants. two pronuclei: one derived from the egg and one from the
wobble position Position 34 in a tRNA, 50 -adjacent to the sperm, After nuclear fusion, it divides mitotically to form the
anticodon. The nucleotide at the position is responsible for embryo.
non-Watson–Crick (wobble) base-pairing.

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