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 The Facts On File

DICTIONARY
of
BOTANY
 The Facts On File

DICTIONARY
of
BOTANY

Edited by
Jill Bailey
The Facts On File Dictionary of Botany

Copyright © 2003 by Market House Books Ltd

All rights reserved. No part of this book may be reproduced or utilized in any
form or by any means, electronic or mechanical, including photocopying,
recording, or by any information storage or retrieval systems, without
permission in writing from the publisher. For information contact:

Checkmark Books
An imprint of Facts On File, Inc.
132 West 31st Street
New York NY 10001

Library of Congress Cataloging-in-Publication Data

The Facts on File dictionary of botany / edited by Jill Bailey.

p. cm.
Includes bibliographical references (p. ).
ISBN 0-8160-4910-6 (hc)—ISBN 0-8160-4911-4 (pbk.)
1. Botany—Dictionaries. I. Title: Dictionary of botany. II. Bailey, Jill.
III. Facts on File, Inc.

QK9.F33 2002
580'.3—dc2l 2002035202

Checkmark Books are available at special discounts when purchased in bulk

quantities for businesses, associations, institutions, or sales promotions. Please call
our Special Sales Department in New York at (212) 967-8800 or (800) 322-8755.

You can find Facts On File on the World Wide Web at

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Compiled and typeset by Market House Books Ltd, Aylesbury, UK

Printed in the United States of America

M P 1 09 8 7 6 5 4 3 2 1

This book is printed on acid-free paper

 PREFACE

This dictionary is one of a series covering the terminology and concepts used
in important branches of science. The Facts On File Dictionary of Botany is
planned as an additional source of information for students taking Ad-
vanced Placement (AP) Science courses in high schools, but will also be help-
ful to older students taking introductory college courses.

This volume covers the whole area of pure and applied plant science includ-
ing anatomy and plant morphology, plant physiology, biochemistry, cell bi-
ology, genetics, evolution, and ecology. It also covers the taxonomy and
classification of plants, with entries for the higher-ranking taxa. The defini-
tions are intended to be clear and informative and, where possible, we have
provided helpful diagrams and examples. The book also has a selection of
short biographical entries for people who have made important contribu-
tions to the field. The appendices include lists of webpages and an informa-
tive bibliography.

The book will be a helpful additional source of information for anyone

studying AP Biology, notably the sections on Organisms and Populations,
Structure and Function of Plants and Animals, and Molecules and Cells.
However, we have not restricted the content to this syllabus. Modern plant
science is a subject of considerable importance and we hope that this book
will be useful to anyone interested in the subject.

ACKNOWLEDGMENTS

Consultant
Andrew Lack B.Sc., Pd.D.

Contributors
Eve Daintith B.Sc.
Elizabeth Tootill B.Sc.
 CONTENTS

Entries A to Z 1

Appendixes

I. SI Units 248

II. Webpages 249

Bibliography 250
 A

ABA See abscisic acid. abscission The organized loss of part of

a plant, usually a leaf, fruit, or unfertilized
abaxial In structures such as a leaves flower. An abscission zone occurs at the
and petals, the side facing away from the base of the organ. Here a separation layer
main axis, i.e. the lower surface. In lateral (abscission layer) is formed by breakdown
organs such as leaves, abaxial is synony- or separation of cells and final severance
mous with the underside. Compare adax- occurs when the vascular bundles are bro-
ial. ken mechanically, e.g. by wind or rain. The
abscission layer is activated by increasing
abiotic environment The nonliving levels of ethylene and a decreasing concen-
factors of the environment that influence tration of auxin
ecological systems. Abiotic factors include
climate, chemical pollution, geographical absolute humidity See humidity.
features, etc.
absolute pollen frequency (APF) See
pollen analysis.

absorption 1. The uptake of liquid by

cells and organs. In plants, water and min-
eral salts are absorbed mainly by the root
hairs, just behind the root tips.
2. The capture of radiant energy by plant
pigments. About 80% of the visible light
falling on a leaf is absorbed, and about
Abscisic acid
10% of the infrared radiation.

abscisic acid (ABA) A plant hormone, absorption spectrum A plot of the ab-
that functions chiefly as an inhibitor of sorbance by a substance of radiation at dif-
growth and cell elongation. Abscisic acid ferent wavelengths, usually of ultraviolet,
has a variety of effects related to seed dor- visible, or infrared radiation. It can give in-
mancy and stress responses: it regulates formation about the identity or quantity of
protein expression in seed development a substance. Chlorophylls, for example,
leading to dormancy and is one of the hor- have absorption peaks in the red and blue
mones involved in bud dormancy; it regu- (and therefore reflect green light). Com-
lates stress responses by, for example, pare action spectrum.
closing stomata in times of water shortage
and increasing the ability of roots to carry accessory cell See subsidiary cell.
water. It can also promote root growth and
inhibit shoot growth. Despite its name it accessory chromosome See B chromo-
does not directly promote abscission, but some.
only indirectly through increasing ethylene
production. Formerly it was known as ab- accessory pigment See photosynthetic
scisin II or dormin. pigments.

1
acellular

acellular Denoting relatively large tis- achene A dry indehiscent fruit formed
sues or organisms that are not composed of from an ovary with a single carpel contain-
discrete cells and are, in effect, unicellular. ing a single seed, e.g. oak (Quercus). Dif-
In flowering plants, for instance, the early ferent types of achenes include the
stages of endosperm development are often CARYOPSIS, CYPSELA, NUT, and SAMARA.
acellular. Other examples include aseptate
fungal hyphae and certain green algae, acicular Needle-shaped.
such as Acetabularia. The term is used in
preference to unicellular to distinguish acid A substance that gives rise to hy-
such structures (which are often multinu- drogen ions (or H3O+) when dissolved in
cleate) from conventional cells and show water. An acid in aqueous solution will
their equivalence to multicellular struc- have a pH below 7. Lowry–Brønsted
tures. See also coenocyte. theory defines an acid as a substance that
exhibits a tendency to release a proton, and
Aceraceae A family of temperate and a base as a substance that tends to accept a
tropical trees and shrubs that includes the proton. Strong acids (e.g. HNO3) react
maples and sycamores. completely with water to give H3O+. Weak
acids (e.g. CH3COOH) are only partly dis-
acetaldehyde (ethanal) An aldehyde, sociated because H3O+ is a stronger acid
CH3CHO, that is an intermediate in the than the free acids.
conversion of pyruvic acid to ethanol dur-
ing the final stage of glycolysis during acidic stain See staining.
anaerobic respiration in plants. It is in-
volved in the synthesis and breakdown of
acid rain The deposition of acids by nat-
ural precipitation, mainly by rain but also
the amino acid threonine.
by snow and fog. Acids are formed by re-
action of gaseous waste products, particu-
acetic acid (ethanoic acid) A carboxylic
larly sulfur dioxide, and also nitrogen
acid, CH3COOH, obtained by the oxida-
oxides, with moisture in the air to form sul-
tion of ethyl alcohol. Acetic acid is a com-
furic and nitric acids. The subsequent pre-
ponent of vinegar (which is obtained by
cipitation has led to raised acidity in forests
bacterial oxidation of wine waste). It is and lakes, especially in Scandinavia and
used as an alternative carbon source by cer- some other parts of northern Europe, dam-
tain green algae. When combined with aging the environment. Acid rain is often
COENZYME A to form ACETYL COA, it plays a
used more loosely for any atmospheric pol-
key role in AEROBIC RESPIRATION. lutant that dissolves in precipitation caus-
ing environmental damage, such as carbon
acetocarmine A stain used to color monoxide or ozone, and these may interact
chromosomes deep reddish-black for view- with the acids.
ing with a light microscope. Tissues are The most serious pollutant is sulfur
fixed in acetic acid before applying aceto- dioxide, which comes mainly from burning
carmine. See fixation; staining. coal and is, consequently, less serious now
than in the twentieth century. Sulfuric acid
acetyl CoA (acetyl coenzyme A) A com- in soils may lead to the formation of am-
pound made up of acetyl and coenzyme A monium sulfate, which causes the release
linked by a sulfur bridge. Acetyl CoA plays of toxic aluminum and heavy metal ions
a key role in metabolism, being a precursor that inhibit metabolic activity. In water-
of the KREBS CYCLE and GLYOXYLATE CYCLE, ways these can damage the gills of fish. Un-
and the starting point for synthesis of fatty polluted rain is normally slightly acidic
acids, terpenes, and some amino acids. The with a pH of 5.0–5.6.
synthesis of acetyl CoA is a high-energy
process, requiring energy from ATP, which acid soil A soil with a pH less than 6.0.
is converted to AMP. Such soils usually form in areas of heavy

2
 activation energy

rainfall, which causes leaching of lime form bacteria are rodlike structures. The
from the surface layers, or over acid sub- Actinobacteria are distinguished by pro-
strata such as granite or sand. In the acidic ducing actinospores: entire cells encysted
conditions, decomposition of organic ma- in thick walls to form resistant spores.
terial in the soil is slow. Acid soils often
contain substantial concentrations of iron actinodromous (palmate; digitate) De-
and aluminum hydroxides. See brown scribing a form of leaf VENATION in which
earth; podsol. three or more primary veins radiate from
the base of the lamina toward the margin,
Acrasiomycota (cellular slime molds) A as in sycamore (Acer pseudoplatanus), re-
class of protoctists in the phylum Rhi- sulting in a leaf that has several large lobes
zopoda. Formerly considered to be fungi, or several leaflets all originating at the
they are made up of independently living same point.
amebas that feed on bacteria, which they
ingest by phagocytosis. Under conditions actinomorphy See radial symmetry.
of food shortage or other stresses, they
come together to form a slug-like aggrega- actinomycetes See Actinobacteria.
tion of amebas called a pseudoplasmo-
dium, which acts like an individual actinostele A type of protostele in
organism. which the xylem is star shaped and the
phloem lies between the points of the
acrocarpous Describing mosses in star. Protosteles are found in the primary
which the reproductive organs are borne at roots of higher plants, and in some spe-
the top of the main axis, which is usually cies of Lycopodium and Psilotum. See
erect, so that subsequent growth is sympo- stele.
dial.
action spectrum A graph showing the
acrocentric See centromere. effect of different wavelengths of radiation,
usually light, on a given process. It is often
acropetal Describing a process (such as similar to the absorption spectrum of the
growth or development) that progresses substance that absorbs the radiation and
from the base or point of attachment, so can therefore be helpful in identifying that
that the oldest parts are at the base and the substance. For example, the action spec-
youngest are at the tip. Compare basipetal. trum of photosynthesis is similar to the ab-
sorption spectrum of chlorophyll. It shows
actin A globular contractile protein (G- the net assimilation plotted against wave-
actin) that makes up 10–15% of the total length. See absorption spectrum. See illus-
cell protein in eukaryotic cells. It is a major tration overleaf.
component of the CYTOSKELETON. It also
controls motility, cellular movements, activation energy The minimum extra
and cyclosis. G-actin can polymerize into energy that must be put into a system to en-
helical strands that coil together to form able a reaction to occur. It increases the en-
microfilaments called fibrous actin (F- ergy levels of participating molecules, and
actin). hence raises their reactivity, often by rais-
ing electrons to an excited state. ENZYMES
Actinobacteria (ray fungi; actinomycetes) enhance reaction rates by lowering the ac-
A phylum of Eubacteria that contains the tivation energy. This is often achieved by
true actinobacteria and the coryneform the attraction of the reactants to the en-
bacteria. The true actinobacteria form zyme’s active site, thus coming into closer
branching filaments that resemble small proximity, or by the enzyme causing con-
fungal mycelia, with hyphae usually less formational or electrostatic changes in one
than 1.5 m m in diameter; the coryne- or more of the reactants.

3
activator

Action spectrum of photosynthesis

Absorption spectrum of chlorophylls

activator 1. A metal ion that acts to- the efficiency of the enzyme activity. See al-
gether with an enzyme or its substrate to losteric site.
bring about a reaction.
2. A protein that positively regulates the active transport The transport of mol-
transcription of a gene. ecules or ions across a cell membrane
against a concentration gradient, with the
active site The particular part of an en- expenditure of energy, usually in the form
zyme molecule that combines with and acts of ATP. It is probably an attribute of all
on the substrate. The active site consists of cells. Anything that interferes with the pro-
amino acids arranged in a configuration vision of energy will interfere with active
specific to a particular substrate or type of transport. The mechanism typically in-
substrate. Binding of an inhibiting com- volves a carrier protein that spans the cell
pound elsewhere on the enzyme molecule membrane and transfers substances in or
may change this configuration and hence out of the cell by changing shape.

4
 ADP

acuminate Describing a structure that Hedyotis on the Hawaiian Islands, but the
gradually narrows to a point, such as cer- term can apply to the range of species in
tain leaves. any genus or family.

acylglycerol (glyceride) An ester of adaxial In structures such as a leaves

glycerol and one or more fatty acids. They and petals, the side facing toward the main
may be mono-, di-, or triacylglycerols axis, i.e. the upper surface. Compare abax-
according to the number of –OH groups ial.
esterified. The fat stores of the body consist
mainly of triacylglycerols (triglycerides). adder’s tongue ferns See Ophioglos-
These can form a source of energy when sales.
carbohydrate levels are low, being broken
down by lipases into fatty acids, which adenine A nitrogenous base (6-amino-
can enter metabolic pathways. Glyco- purine) found in DNA and RNA. It is also
lipids (glycosyldiacylglycerols) are diacyl- a constituent of certain coenzymes, e.g.
glycerols with a sugar attached to the unes-
NAD and FAD, and when combined with
terified hydroxyl group. See also lipid.
the sugar ribose it forms the nucleoside
ADENOSINE found in AMP, ADP, and ATP.
adaptation The extent to which an or-
Adenine has a purine ring structure.
ganism, or a physiological or structural
characteristic of an organism, is suited to a
particular environment. Specialist organ- adenosine (adenine nucleoside) A nu-
isms that have become highly adapted to cleoside formed from adenine linked to D-
one environment are then often not so ribose with a b-glycosidic bond. Adenosine
adaptable as less specialized organisms triphosphate (ATP) is a nucleotide derived
and are at a disadvantage in a changing en- from adenosine.
vironment (adaptation versus adaptabil-
ity). adenosine diphosphate See ADP.

adaptive enzyme See inducible enzyme. adenosine monophosphate See AMP.

adaptive radiation The formation adenosine triphosphate See ATP.

through evolution of a number of different
varieties or species from a common ances- Adiantaceae (maidenhair ferns) A fam-
tor. It is often seen most clearly on isolated ily of ferns (Filicinophyta) found through-
oceanic islands, e.g. the many forms of out the world, especially in the moist
trees, shrubs, vines, and herbs of the genus American tropics.

Active site

5
adventitious

Aerobic respiration

ADP (adenosine diphosphate) A nu- aerobe An organism that can live and
cleotide consisting of adenine and ribose grow only in the presence of free oxygen,
with two phosphate groups attached. The i.e. it respires aerobically (see aerobic res-
second phosphate is attached to the first by piration). All plants and most bacteria and
a high-energy bond. fungi are aerobes. Compare anaerobe.

adventitious Describing plant organs aerobic respiration Respiration in

that arise in unexpected places, for exam- which free oxygen is used to oxidize or-
ple the development of adventitious roots ganic substrates to carbon dioxide and
from stems, and adventitious buds from water, with a high yield of energy. The re-
leaves. action overall is:
C6H12O6 + 6O2 = 6CO2 + 6H2O +
aerenchyma A plant tissue containing energy
It occurs in a number of stages, the first of
large intercellular air spaces, usually
which (GLYCOLYSIS) also occurs in anaero-
formed as a consequence of the death of
bic respiration in the cell cytoplasm. With
parenchyma cells. Aerenchyma is typical of
glucose as the substrate, a sequence of re-
the stems of many aquatic plants whose
actions results in the formation of pyru-
leaves float at the surface of ponds and vate. The remaining stages, which do not
lakes, giving added buoyancy to the tis- occur in anaerobic respiration, take place
sues. in the mitochondria. Pyruvate is converted
to ACETYL COA, which enters a cyclic series
aerial root A root that arises above soil of reactions, the KREBS CYCLE, with the pro-
level. Examples include the roots of epi- duction of carbon dioxide and hydrogen
phytes and climbers, which hang down in atoms. These and other hydrogen atoms
the air or stick to a trunk or branch. The produced at earlier stages are now passed
roots of many orchids and other epiphytes to the ELECTRON-TRANSPORT CHAIN (involv-
developed a sheath of dead cells, the vela- ing CYTOCHROMES and FLAVOPROTEINS).
men, which helps to absorb water from the Here they combine with atoms of free oxy-
atmosphere. gen to form water. Energy released at each

6
 air bladder

stage of the chain is used to form ATP dur- gels are extensively used for growing
ing a coupling process (see oxidative phos- microorganisms and tissue cultures.
phorylation). There is a net production of
38 ATPs (2930 kJ) per molecule of glucose Agaricales (agarics) An order of basid-
during aerobic respiration, a yield of about iomycetes that contains the mushrooms
19 times that of anaerobic respiration (300 and toadstools. The cap of the fleshy fruit-
kJ), and the mechanism of the majority of ing body – the mushroom or toadstool –
organisms. Compare anaerobic respira- bears a series of parallel gills on its lower
tion. surface that greatly increase its surface
area. These are covered in the spore-
aerotaxis (aerotactic movement) A bearing layer (hymenium). Agarics are
TAXIS in response to an oxygen concentra- common in leaf litter and rotting wood,
tion gradient. For instance, motile aerobic where they live as saprobes.
bacteria are positively aerotactic, whereas
motile obligate anaerobic bacteria are neg- aggregate fruit (compound fruit) A fruit-
atively aerotactic. Some photosynthetic eu- like structure that is made up of several in-
karyotic unicells are also aerotactic. dividual fruits derived from the carpels of a
single flower, e.g. strawberry (Fragaria),
aerotropism A TROPISM in which the blackberry (Rubus fruticosus). These may
orientating stimulus is oxygen. be achenes, follicles, berries, or drupelets.
An aggregate fruit is sometimes called an
afforestation The establishment of for- etaerio, but this term may also be restricted
est on land not previously forested, either to an aggregate of drupelets. A fruit that
by natural succession or by planting. develops from a group of flowers is termed
a multiple fruit, e.g. pineapple (Ananas co-
aflatoxin One of a group of carcino- mosus) and hop (Humulus lupulus).
genic toxins produced by fungi of the genus
Aspergillus, especially A. flavus. This Agrobacterium A genus of soil bacte-
species is a common contaminant of crops, ria, the species A. tumefaciens being the
such as peanuts. Contaminated feed causes causative agent of CROWN GALL, a type of
serious outbreaks of disease among live- tumor in plants. A segment of DNA (trans-
stock. ferred DNA, T-DNA) from a PLASMID in
the bacterium is transferred into the host
after-ripening The collective name for DNA and induces tumor formation. Since
processes that are necessary before germi- the plasmid is capable of independent
nation can take place in certain seeds, even replication in host cells of many dicotyle-
though external conditions may be suit- donous plants, it has been used as a cloning
able. For example, a period of dormancy vector in GENETIC ENGINEERING.
may be imposed on the seed, preventing
premature germination before an unfavor- agroforestry A system of cultivation
able season such as winter. After-ripening common in many parts of the tropics, es-
is common in areas with marked seasonal- pecially in rainforest regions, in which
ity. forestry and arable farming are mixed.
This reduces soil degradation, as freshly
agamospermy See apomixis. tilled soils are shaded and protected from
heavy rains and run-off by the trees. The
agar A gelling agent prepared from sea- trees help to draw nutrients up from the
weed (normally a red alga, Gelidium), used deeper soil layers into the zone occupied by
to set liquid nutrients. It can withstand the roots of the crop plants.
sterilization at high temperatures and is re-
sistant to attack by most bacteria. Agar agronomy A branch of agriculture deal-

7
air plant

ing with soil management and crop pro- alcohol dehydrogenase An enzyme
duction. that converts ethanol into acetaldehyde
(ethanal). It is important in a plant’s ability
air bladder 1. A structure on the pollen to withstand waterlogging. Ethanol is a
grains of some conifers, such as Pinus, that product of anaerobic respiration in plants,
aids the wind dispersal of the pollen. It and waterlogged roots suffer from a short-
consists of an air-filled protuberance on age of oxygen. Accumulation of ethanol is
each side of the pollen grain developed by harmful to a plant.
the separation of the exine layers, giving
the pollen a characteristic winged appear- alcoholic fermentation A form of
ance. ANAEROBIC RESPIRATION in which glucose is
2. An air-filled chamber in the thallus of broken down to form ethanol and carbon
many Phaeophyta (brown algae) that in- dioxide. It is carried out by yeasts and
creases the buoyancy of the thallus, en- some other fungi and certain bacteria.
abling it to float toward the surface when Alcoholic fermentation is catalyzed by
underwater to increase light absorption enzymes of the zymase complex, which are
and gaseous exchange for photosynthesis. secreted by the cells or released after the
Air bladders are highly developed in cer- cells die. The process is self-limiting, since
tain wracks, such as the bladder wrack the organisms usually die once the alcohol
(Fucus vesiculosus). concentration in the medium exceeds 15%.
Alcoholic fermentation is the basis of the
air plant See epiphyte. preparation of alcoholic beverages, and of
bread making (in which the carbon dioxide
alanine A simple AMINO ACID, one of the
early products of photosynthesis. Alanine
is formed by transamination when an
amino group is donated by gutamine to
pyruvic acid. It may be deaminated back to
pyruvate for use in the KREBS CYCLE.

albumin One of a group of simple, low-

molecular-weight proteins found in plants,
for example in the endosperm of barley
and wheat seeds. Albumins are water-
soluble and coagulate when heated.

albuminous cell 1. A vertically elon-

gated parenchyma cell, found in groups in
the rays of the secondary phloem in gym-
nosperms, where they are associated with
sieve cells. Unlike the companion cells of
angiosperms, they are not derived from the
same mother cell as the sieve cell.
2. An albumin-containing cell found in cer-
tain seeds.

alcohol A type of organic compound of

the general formula ROH, where R is a hy-
drocarbon group. Examples of simple alco-
hols are methanol (CH3OH) and ethanol
(C2H5OH). Ethanol is a product of anaer-
obic metabolism.
Alcoholic fermentation

8
 alkaloid

released causes the dough to rise). During carbon atoms) and glucose (with six car-
alcoholic fermentation acetaldehyde acts bon atoms).
as a hydrogen acceptor instead of oxygen.
The pyruvic acid formed by GLYCOLYSIS is aleurone grain (aleurone body) A mod-
broken down into acetaldehyde and car- ified vacuole found in the embryo and en-
bon dioxide. The acetaldehyde is further dosperm of seeds and containing mostly
reduced by NADH to form ethanal, releas- reserve proteins, but also phytic acid and
ing only about a tenth of the energy that various enzymes associated with mobiliza-
would be released by aerobic respiration. tion (digestion) of these reserves. The pro-
teins and phytic acid are present in
aldehyde A type of organic compound crystalline form in the dormant seed.
with the general formula RCHO, where
the –CHO group (the aldehyde group) con- aleurone layer The outermost protein-
sists of a carbonyl group attached to a hy- rich layer of the endosperm of grass fruits
drogen atom. Simple examples of (e.g. cereal grains). At germination, the em-
bryo produces gibberellin, which stimu-
lates the aleurone layer to synthesize
enzymes, especially amylase. The latter
causes hydrolysis of the starch in the en-
dosperm. The enzymes are synthesized
from the amino acids supplied by break-
down of ALEURONE GRAINs.

algae (sing. alga) A large mixed group of

photosynthesizing eukaryotic organisms,
now usually placed in the kingdom Protoc-
tista. They often resemble plants and are
found mainly in marine or fresh-water
habitats, although some algae are terres-
trial. Algae differ from plants in lacking
any real differentiation of leaves, stems,
and roots, in having no layer of sterile cells
around the reproductive organs,and in not
having an embryo stage in their life cycle.
Algae can be unicellular (e.g. Chlamy-
domonas), colonial (e.g. Volvox), filamen-
tous (e.g. Spirogyra), or thalloid (e.g.
Fucus). All algae contain chlorophyll but
this may be masked by various accessory
pigments, these being one of the major
characteristics used to divide the algae
into their various phyla. Other characters
used to classify the algae are the nature
Aldose
of storage products, the type of cell wall,
the form and number of undulipodia
aldehydes are methanal (formaldehyde, (flagella), ultrastructural cell details, type
HCHO) and ethanal (acetaldehyde, of food reserves, and reproductive pro-
CH3CHO). cesses.

aldose A SUGAR containing an aldehyde alkaline soil See calcareous soil.

(CHO) or potential aldehyde group. Ex-
amples include the sugars ribose (with five alkaloid One of a group of organic com-

9
allele

pounds found in plants, which are poiso- DIOECIOUSand MONOECIOUS. Most plants
nous insoluble crystalline compounds. are hermaphrodite, but have mechan-
They contain nitrogen and usually occur as isms such as self-INCOMPATIBILITY that
salts of acids such as citric, malic, and suc- promote allogamy. Compare autogamy.
cinic acids. Their function in plants re- See also heterostyly; chasmogamy; di-
mains obscure, but it is suggested that they chogamy.
may be nitrogenous end-products of me-
tabolism, or they may have a protective allopatric species See species.
function against herbivores, since they
taste bitter. Important examples in human allopolyploidy A type of polyploidy in-
use are quinine, nicotine, atropine, opium, volving the combination of chromosomes
morphine, codeine, and strychnine. They from two or more different species. Al-
occur mainly in the poppy family, the but- lopolyploids usually arise from the dou-
tercup family, and the nightshade family of bling of chromosomes of a hybrid between
plants. species, the doubling often making the hy-
brid fertile. The properties of the hybrid,
allele (allelomorph) One of the possible such as greater vigor and adaptability, are
forms of a given gene. The alleles of a par- retained in the allopolyploid in subsequent
ticular gene occupy the same positions generations and such organisms are often
(loci) on homologous chromosomes. A highly successful. Many plant species have
gene is said to be homozygous if the two been derived originally from allopoly-
loci have identical alleles and heterozygous
ploidy, e.g. cultivated wheat. Compare
when the alleles are different. When two
autopolyploidy.
different alleles are present, one (the domi-
nant allele) usually masks the effect of the
allosteric enzyme An enzyme whose
other (the recessive allele). The allele deter-
catalytic activity can be modified by the
mining the normal form of the gene is usu-
noncovalent binding of a particular
ally dominant while mutant alleles are
metabolite (a modulator) at a site (the AL-
usually recessive. Thus most mutations
LOSTERIC SITE) other than the active site. It
show in the phenotype only when they are
homozygous. In some cases one allele is not may either inhibit or enhance the enzyme
completely dominant or recessive to an- activity. Allosteric enzymes may have more
other allele (INCOMPLETE DOMINANCE). than one modulator.
Thus an intermediate phenotype will be
produced in the heterozygote. See also in- allosteric site A part of an enzyme to
complete dominance; multiple allelism. which a specific effector or modulator can
be attached. This attachment is reversible.
allelomorph See allele. Allosteric enzymes possess an allosteric site
in addition to their ACTIVE SITE.
allelopathy Inhibition of the germina-
tion, growth, or reproduction of an organ- allotetraploid (amphidiploid) An allo-
ism effected by a chemical substance polyploid whose chromosomes are derived
released from another organism. It is prob- from two different species and which
ably an anticompetition mechanism in therefore has four times the haploid num-
plants; for example, barley secretes an al- ber of chromosomes, e.g. Spartina anglica,
kaloid substance from its roots to inhibit derived from the diploids S. alterniflora
competing weeds, but it is not known how and S. maritima. See allopolyploidy.
important it is generally.
alluvial soil A type of soil formed on
allogamy Cross-fertilization in plants. river floodplains and deltas, where new
This promotes genetic variation in the sediment is deposited on the land during
population especially in plants that are floods.

10
 amino acids

alpha helix A highly stable structure in alternation of generations The occur-

which peptide chains are coiled to form a rence of two, or occasionally more, gener-
spiral. Each turn of the spiral contains ap- ations during the life cycle of an organism.
proximately 3.6 amino acid residues. The In all plants and some algae there is an al-
R group of these amino acids extends out- ternation between sexual haploid and
ward from the helix. Hydrogen bonding asexual diploid stages. They usually differ
between successive coils holds the helix to- markedly in morphology. The haploid
gether. If the alpha helix is stretched the plant produces gametes mitotically and is
hydrogen bonds are broken but reform on thus termed the gametophyte while the
relaxation. The alpha helix is found in diploid plant produces spores meiotically
muscle protein and keratin. and is called the sporophyte. The gametes
fuse to form a zygote, which develops into
alpha-naphthol test (Molisch’s test) A the sporophyte, and the spores germinate
test for detecting the presence of carbohy- and produce the gametophyte, so forming
drates in solution. The test solution is a cycle. In bryophytes (mosses, liverworts
placed in a test tube, and a small amount of and hornworts) the haploid gametophyte is
alpha-naphthol added; concentrated sulfu- the dominant phase of the life cycle and the
ric acid is then trickled slowly down the sporophyte is represented only by the cap-
side of the tube. If carbohydrate is present, sule, seta, and foot. In vascular plants the
a violet ring will form at the junction of the diploid sporophyte is the dominant phase
liquids. and in the ferns, for example, the gameto-
phyte is a small prothallus. The concept of
alpine Describing a BIOME (regional an alternation of generations can be ex-
community) of plants above the treeline tended to the flowering plants, in which the
and below the snowline on high moun- embryo sac and pollen represent the much
tains. The lower limit of the alpine zone reduced female and male gametophyte gen-
varies in different mountain regions, ac- erations respectively.
cording to the rainfall and other climatic
and topographic factors, from 100 meters ameboid Describing an organism that
above sea level in parts of Scotland to 3700 resembles an ameba in shape and move-
meters in the western Himalayas. There are ment.
often considerable differences between the
nature and distribution of vegetation on amensalism An association between
north- and south-facing slopes and be- two different species, at either the level of
tween windward and leeward slopes. the individual organism or the population
Alpine vegetation is often similar to TUN- level, in which one is harmed and the other
DRA vegetation, being adapted to harsh cli- is unaffected. Compare commensalism;
mate conditions, including high wind mutualism.
speeds.
amino acids Compounds containing
alternate Describing a leaf arrangement both carboxylic acid and amino groups in
in which there is only one leaf at each node, their molecules. The amino acids have the
as in hazel. This is the commonest form of general formula RCHNH2–COOH. Here,
leaf arrangement. See phyllotaxis. the group R ranges from a simple hydrogen
atom to complex ring structures. All amino
alternate host Any host other than acids are white, crystalline, soluble in
the main (most common) one. For exam- water, and with the sole exception of the
ple, many rusts overwinter on alternate simplest member, glycine, all are optically
hosts. Some parasites and pests are able to active.
live on more than one host, and some Amino acids are the basic components
need a second host to complete their life of proteins, which consist of chains of
cycle. amino acids. Plants also contain some 200

11
amino acid sequencing

Amino acids

amino acids that do not occur in proteins, Amino acids contain both acids (car-
some of which have protective or storage boxylic acids) and bases (amino group), so
roles, while others are intermediates in the they react with both acids and bases. Thus
synthesis of commoner amino acids. the charge on the amino acid depends on

12
 amitosis

Amino acids

the pH. This feature is used to separate

amino acids for analysis by chromatogra-
phy and other methods. Each amino acid
has its own characteristic isoelectric point,
where the net charge on the molecule is
zero. See illustration overleaf.

13
AMP

amino acid sequencing The determina- the starch molecule, while a-amylase can
tion of the amino acid sequence of a pro- attack bonds within the starch molecule.
tein or peptide. This sequence is the primary Both a- and b-amylases occur in plants, the
structure of a protein; it influences sec- latter particularly in malt (being used in the
ondary, tertiary, and quaternary structure. brewing industry), but only b-amylase is
found in animals, having an important role
amino sugar A monosaccharaide sugar in digestion. During seed germination a-
in which a hydroxyl group (OH) has been amylase breaks down the starch of the
replaced by an amino group (NH2). Glu- endosperm, the transcription of the a-
cosamine (from glucose) is one of the amylase gene being greatly enhanced by
commonest amino sugars. Galactosamine gibberellic acid produced by the embryo.
(from galactose) is a major component of a-amylase is also involved in the release of
glycolipids. Amino sugars are important stored starch from plastids.
components of bacterial cell walls.
amylopectin The water-insoluble frac-
amitosis Nuclear division characterized tion of STARCH. It is a branching polymer of
by the absence of a nuclear spindle and glucose units.
leading to the production of daughter nu-
clei with unequal sets of chromosomes. amyloplast A plastid that synthesizes
The ordered process of division, duplica- and stores starch grains. Amyloplasts are
tion of chromosomes, dissolution of nu- common in storage organs, e.g. the potato
clear membrane, and production of a tuber. They have a physiological role in the
spindle as in mitosis is apparently absent. root cap and elsewhere, where the starch
Cells produced amitotically inherit vari- grains act as statoliths.
able numbers of chromosomes. The
chances of a daughter cell lacking essential amylose The water-soluble fraction of
genes are less than may be expected since STARCH. It is an unbranched polymer of
many cells that characteristically divide glucose units.
amitotically are polyploid, e.g. the en-
dosperm nucleus in angiosperms. Compare anabolism Metabolic reactions in
endomitosis; mitosis. which molecules are linked together to
form more complex compounds. Thus, an-
AMP (adenosine monophosphate) A abolic reactions are concerned with build-
nucleotide consisting of adenine, ribose, ing up structures, storage compounds, and
and phosphate. AMP has an important complex metabolites in the cell. Starch,
role in the regulation of glycolysis, pro- fats, and proteins are all products of ana-
moting the production of fructose bisphos- bolic pathways. Anabolic reactions gener-
phate from fructose 6-phosphate. See ATP; ally require an input of energy, usually
cyclic AMP. provided by ATP produced by catabolism.
An example is the Calvin cycle. See also
amphidiploid See allotetraploid. metabolism. Compare catabolism.

amphimixis True sexual reproduction anaerobe An organism that can live and
by fusion of gametes. Compare apomixis. grow in the absence of free oxygen, i.e. it
respires anaerobically (see anaerobic respi-
amylase An enzyme, found widely in ration). Anaerobes can be facultative, in
plants, animals, and microorganisms, that that they usually respire aerobically but
hydrolyzes starch or glycogen to the sugars can switch to anaerobic respiration when
maltose, glucose, or dextrin. Amylase hy- free oxygen is in short supply, as are most
drolyzes the a(1-4)glycosidic bonds in yeasts; or obligate, in that they never
starch. It occurs in two forms, a and b. b- respire aerobically and may even be poi-
amylase attacks the nonreducing ends of soned by free oxygen, as are denitrifying

14
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