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E

e The irrational number dened as the *broken symmetries in *gauge theories


limit as n tends to innity of (1 + 1/n)n. It become unbroken symmetries. As the
has the value 2.718 28. It is used as the universe cools after the big bang there is
base of natural *logarithms and occurs in thought to be a sequence of transitions to
the *exponential function, ex. broken symmetry states.
Combining cosmology with grand
ear The sense organ in vertebrates that unied theories helps to explain why the
is specialized for the detection of sound observed universe appears to consist of
and the maintenance of balance. It can be matter with no antimatter. This means
divided into the *outer ear and *middle that one has a nonzero *baryon number
ear, which collect and transmit sound for the universe. This solution relies on
waves, and the *inner ear, which con- the fact that there were nonequilibrium
tains the organs of balance and (except in conditions in the early universe due to its
sh) hearing (see illustration). rapid expansion after the big bang.
eardrum See tympanum. An important idea in the theory of the
early universe is that of ination the
early universe The study of *cosmol- idea that the nature of the *vacuum state
ogy at the time very soon after the *big gave rise, after the big bang, to an expo-
bang. Theories of the early universe have nential expansion of the universe. The
led to a mutually benecial interaction hypothesis of the inationary universe
between cosmology and the theory of *el- solves several long-standing problems in
ementary particles, particularly *grand cosmology, such as the atness and
unied theories. homogeneity of the universe. Quantum
Because there were very high tempera- uctuations during ination have been
tures in the early universe many of the suggested as a mechanism for the forma-

semicircular
canals utriculus

ear ossicles
pinna perilymph
malleus stapes
(hammer) (stirrup)
incus ampulla
(anvil)
auditory nerve

sacculus
oval window
nerve fibre
tympanum
(eardrum) cochlea

external endolymph
auditory meatus round
window Eustachian tube

middle
outer ear ear inner ear

Structure of the mammalian ear


263 earths atmosphere

tion of large-scale structures. Evidence for focus; the point on the earths surface di-
this theory has been provided by *COBE rectly above this is the epicentre (or
and by *WMAP. hypocentre). See seismic waves. Earth-
quakes result from a build-up of stresses
ear ossicles Three small bones the
within the rocks until they are strained to
incus (anvil), malleus (hammer), and stapes
the point beyond which they will fracture.
(stirrup) that lie in the mammalian
They occur in narrow continuous belts of
*middle ear, forming a bridge between
activity, which correspond with the junc-
the tympanum (eardrum) and the *oval
tion of lithospheric plates, including the
window. The function of the ossicles is to
transmit (and amplify) vibrations of the
circum-Pacic belt, the AlpineHimalayan e
belt, and mid-ocean ridges. The scale of
tympanum across the middle ear to the
the shock of an earthquake is known as
oval window, which transfers them to the
the magnitude; the most commonly used
*inner ear. Muscles of the middle ear con-
scale for comparing the magnitude of
strict the movement of the ossicles. This
earthquakes is the logarithmic *Richter
serves to safeguard the ear from damage
scale (9.5 is the highest recorded magni-
caused by excessively loud noise.
tude on the scale).
earth The planet that orbits the sun be-
earths atmosphere The gas that sur-
tween the planets Venus and Mars at a
rounds the earth. The composition of dry
mean distance from the sun of
air at sea level is: nitrogen 78.08%, oxygen
149 600 000 km. It has a mass of about
20.95%, argon 0.93%, carbon dioxide
5.974 1024 kg and an equatorial diame-
0.03%, neon 0.0018%, helium 0.0005%,
ter of 12 756.3 km. The earth consists of
krypton 0.0001%, and xenon 0.00001%. In
three layers: the gaseous atmosphere (see
addition to water vapour, air in some lo-
earths atmosphere), the liquid *hydro-
calities contains sulphur compounds,
sphere, and the solid *lithosphere. The
hydrogen peroxide, hydrocarbons, and
solid part of the earth also consists of
dust particles.
three layers: the crust with a mean thick-
The lowest level of the atmosphere, in
ness of about 32 km under the land and
which most of the weather occurs, is
10 km under the seas; the mantle, which
called the troposphere. Its thickness
extends some 2900 km below the crust;
varies from about 7 km at the poles to
and the core, part of which is believed to
28 km at the equator and in this layer
be liquid. The crust has a relative density
temperature falls with increasing height.
of about 3 and consists largely of sedimen-
The next layer is the stratosphere, which
tary rocks overlaying igneous rocks. The
goes up to about 50 km. Here the temper-
composition of the crust is: oxygen 47%,
ature remains approximately constant.
silicon 28%, aluminium 8%, iron 4.5%, cal-
Above this is the ionosphere, which ex-
cium 3.5%, sodium and potassium 2.5%
tends to about 1000 km, with the temper-
each, and magnesium 2.2%. Hydrogen,
ature rising and the composition
carbon, phosphorus, and sulphur are all
changing substantially. At about 100 km
present to an extent of less than 1%. The
and above most of the oxygen has dissoci-
mantle reaches a relative density of about
ated into atoms; at above 150 km the per-
5.5 at its maximum depth and is believed
centage of nitrogen has dropped to nil. In
to consist mainly of silicate rocks. The
the ionosphere the gases are ionized by
core is believed to have a maximum
the absorption of solar radiation. This en-
relative density of 13 and a maximum
ables radio transmissions to be made
temperature of 6400 K. See also geo-
round the curved surface of the earth as
magnetism; plate tectonics.
the ionized gas acts as a reector for cer-
earthquake A sudden movement or tain wavelengths. The ionosphere is di-
fracturing within the earths lithosphere, vided into three layers. The D-layer
causing a series of shocks. This may range (5090 km) contains a low concentration
from a mild tremor to a large-scale earth of free electrons and reects low-
movement causing extensive damage over frequency radio waves. The E-layer
a wide area. The point at which the earth- (90150 km) is also called the Heaviside
quake originates is known as the seismic layer or HeavisideKennelly layer as its ex-
earth sciences 264

550

500 exosphere

450

400

e 350 F region

300
km

250 (Appleton
layer)
ionosphere
(HeavisideKennelly
200 layer)

150
E region
100
D region
50
stratosphere
troposphere
0 0 2 4 6 8 10 12 14
log10 density (kg m3)

Earths atmosphere

istence was predicted independently by by earthshine, as things on earth may be


Oliver Heaviside (18501925) and Arthur illuminated by moonlight. Under certain
E. Kennelly (18611939). This layer reects conditions near new moon the dark disc
medium-frequency waves. The F-layer of the moon can be seen faintly illumi-
(1501000 km) is also called the Appleton nated by earthshine a phenomenon
layer after its discoverer Sir Edward Ap- called the old moon in the new moons
pleton (18921965). It has the highest con- arms.
centration of free electrons and is the
earthslide The movement of a layer of
most useful for radio transmission. Wave-
dry soil down a slope. The soil layer is in-
lengths between 8 mm and 20 m are not
herently unstable and shears, sometimes
reected by the ionosphere but escape
brought about by the action of water. But
into space. Therefore television transmis-
if the soil becomes very wet, and the
sions, which utilize this range, require
slope is steep, an earthow may occur.
articial *satellites for reection (or recep-
Contributory factors are sparse vegetation
tion, amplication, and retransmission).
and sudden rainfall.
From about 400 km, the outermost region
of the atmosphere is also called the exo- earths magnetic eld See geomag-
sphere. See illustration. netism.
earth sciences A group of sciences con- earwigs See dermaptera.
cerned with the study of the earth. The
ebonite See vulcanite.
chief earth sciences are geology, physical
geography, oceanography, meteorology, ebullioscopic constant See elevation
geophysics, and geochemistry. of boiling point.
earthshine Sunlight reected from the ebullioscopy The use of *elevation of
surface of the earth. An observer in space boiling point to determine relative mo-
may see nearby objects dimly illuminated lecular masses.
265 ecliptic

eccentricity See conic. in the skin. In many species (e.g. sea


urchins) spines protrude from the test. A
Eccles, Sir John Carew (190397) Aus-
system of water-lled canals (the water
tralian physiologist, who was educated in
vascular system) provides hydraulic power
Melbourne and Oxford, and held appoint-
for thousands of tube feet: saclike pro-
ments in Britain, Australia, New Zealand
trusions of the body wall used for loco-
and, nally, the USA. While in Australia
motion, feeding, and respiration.
he carried out his best-known work, on
Echinoderms have a long history: fossils
the transmission of nerve impulses across
of primitive echinoderms are known from
synapses, which he attributed to the neu-
rotransmitter acetylcholine. He shared
rocks over 500 million years old. e
the 1963 Nobel Prize for physiology or echo The reection of a wave by a sur-
medicine with Sir Alan Hodgkin (191498) face or object so that a weaker version of
and Sir Andrew Huxley (1917 ), who it is detected shortly after the original.
worked in the same area of biology. The delay between the two is an indica-
tion of the distance of the reecting sur-
ecdysis (moulting) 1. The periodic loss
face. An echo sounder is an apparatus for
of the outer cuticle of arthropods. It starts
determining the depth of water under a
with the reabsorption of some materials
ship. The ship sends out a sound wave
in the inner part of the old cuticle and the
and measures the time taken for the echo
formation of a new soft cuticle. The re-
to return after reection by the sea bot-
mains of the old cuticle then split; the
tom. Sonar (sound navigation ranging) is a
animal emerges and absorbs water or
technique for locating underwater objects
swallows air and increases in size while
by a similar method. Echoes also occur
the new cuticle is still soft. This cuticle is
with radio waves; reection of waves
then hardened with chitin and lime salts.
causes an echo in radio transmission and
In insects and crustaceans ecdysis is con-
ghosts in television pictures. See also
trolled by the hormone *ecdysone. 2. The
radar.
periodic shedding of the outer layer of the
epidermis of reptiles (except crocodiles) to echolocation 1. See radar; echo. 2. A
allow growth to occur. method used by some animals (such as
bats, dolphins, and certain birds) to detect
ecdysone A steroid hormone, produced
objects in the dark. The animal emits a se-
by insects and crustaceans, that stimu-
ries of high-pitched sounds that echo back
lates moulting (see ecdysis) and metamor-
from the object and are detected by the
phosis. It acts on specic gene loci,
ear or some other sensory receptor. From
stimulating the synthesis of proteins in-
the direction of the echo and from the
volved in these bodily changes.
time between emission and reception of
ECG See electrocardiogram. the sounds the object is located, often
very accurately.
echelon A form of *interferometer con-
sisting of a stack of glass plates arranged ECL See emitter-coupled logic.
stepwise with a constant offset. It gives a
eclipse The total (total eclipse) or partial
high resolution and is used in spectros-
(partial eclipse) obscuring of light from a
copy to study hyperne line structure. In
celestial body as it passes behind or
the transmission echelon the plates are
through the shadow of another body. A
made equal in optical thickness to intro-
lunar eclipse occurs when the sun, earth,
duce a constant delay between adjacent
and moon are in a straight line and the
parts of the wavefront. The reecting
shadow of the earth falls on the moon. A
echelon has the exposed steps metallized
solar eclipse occurs when the shadow of
and acts like an exaggerated *diffraction
the moon falls on the earth. See illustra-
grating.
tions.
Echinodermata A phylum of marine
eclipsed conformation See conforma-
invertebrates that includes the sea
tion.
urchins, starsh, brittlestars, and sea cu-
cumbers. Echinoderms have an exoskele- ecliptic The *great circle in which the
ton (test) of calcareous plates embedded plane of the earths orbit round the sun
ECM 266

have led to increased concern about the


effects of human activities on the environ-
lunar ment (notably the effects of *pollution),
eclipse
which has resulted in a greater awareness
of the importance of *conservation.
ecosystem A biological *community
earths and the physical environment associated
orbit
with it. Nutrients pass between the differ-
e solar
eclipse
ent organisms in an ecosystem in denite
pathways; for example, nutrients in the
soil are taken up by plants, which are
then eaten by herbivores, which in turn
may be eaten by carnivores (see food
chain). Organisms are classied on the
basis of their position in an ecosystem
Solar and lunar eclipses
into various *trophic levels. Nutrients and
energy move round ecosystems in loops
or cycles (in the case above, for example,
moons nutrients are returned to the soil via ani-
motion mal wastes and decomposition). See car-
bon cycle; nitrogen cycle.
ectoderm The external layer of cells of
the *gastrula, which will develop into the
path of epidermis and the nervous system in the
totality
earths adult. See also germ layers.
motion
ectoparasite A parasite that lives on
Moon's shadow in solar eclipse the outside of its hosts body. See para-
sitism.
intersects the *celestial sphere. It is thus ectoplasm See cytoplasm.
the suns apparent annual path across the
sky. Ectoprocta See bryozoa.

ECM See extracellular matrix. ectotherm (poikilotherm) An animal


that maintains its body temperature by
E. coli See escherichia coli. absorbing heat from the surrounding en-
ecological niche The status or role of vironment. All animals except mammals
an organism in its environment. An or- and birds are ectotherms; they are often
ganisms niche is dened by the types of described as being cold-blooded and are
food it consumes, its predators, tempera- unable to regulate their body temperature
ture tolerances, etc. Two species cannot metabolically. See poikilothermy. Compare
coexist stably if they occupy identical endotherm.
niches. edaphic factor A factor relating to the
ecology The study of the interrelation- physical or chemical composition of the
ships between organisms and their natu- soil found in a particular area. For exam-
ral environment, both living and ple, very alkaline soil may be an edaphic
nonliving. For this purpose, ecologists factor limiting the variety of plants grow-
study organisms in the context of the ing in a region.
*populations and *communities in which Eddington limit A limit for the maxi-
they can be grouped and the *ecosystems mum value of the brightness of a star of a
of which they form a part. The study of given mass. This limit exists because the
ecological interactions provides important radiation pressure caused by the nuclear
information on the nature and mecha- fusion reactions powering the star has to
nisms of evolutionary change. Advances counter, but not exceed, the gravitational
made in ecology over the last 25 years force that would cause gravitational col-
267 egg

lapse of the star. The existence of this from the central nervous system to an *ef-
limit was rst pointed out by the English fector in order to bring about a physiolog-
astrophysicist Sir Arthur Stanley Edding- ical response to changes in the
ton (18821944). environment.
eddy current A current induced in a efferent Carrying (nerve impulses,
conductor situated in a changing mag- blood, etc.) away from the centre of a
netic eld or moving in a xed one. Any body or organ towards peripheral regions.
imagined circuit within the conductor The term is usually applied to types of
will change its magnetic ux linkage, and nerve bres or blood vessels. Compare af-
the consequent induced e.m.f. will drive ferent.
e
current around the circuit. In a substan-
tial block of metal the resistance will be
effervescence The formation of gas
bubbles in a liquid by chemical reaction.
small and the current therefore large.
Eddy currents occur in the cores of trans- efciency A measure of the perfor-
formers and other electrical machines and mance of a machine, engine, etc., being
represent a loss of useful energy (the the ratio of the energy or power it deliv-
eddy-current loss). To reduce this loss to a ers to the energy or power fed to it. In
minimum metal cores are made of insu- general, the efciency of a machine varies
lated sheets of metal, the resistance be- with the conditions under which it oper-
tween these laminations reducing the ates and there is usually a load at which it
current. In high-frequency circuits *ferrite operates with the highest efciency. The
cores can be used. Eddy currents in a mov- thermal efciency of a heat engine is the
ing conductor interact with the magnetic ratio of the work done by the engine to
eld producing them to retard the motion the heat supplied by the fuel. For a re-
of the conductor. This enables some elec- versible heat engine this efciency equals
trical instruments (moving-coil type) to (T1 T2)/T1, where T1 is the thermody-
utilize eddy currents to create damping. namic temperature at which all the heat
Eddy currents are also used in *induction is taken up and T2 is the thermodynamic
heating. temperature at which it is given out (see
Edison cell See nickeliron accumula- carnot cycle). For real engines it is al-
tor. ways less than this.

EDTA Ethylenediaminetetraacetic acid, eforescence The process in which a


crystalline hydrate loses water, forming a
(HOOCCH2)2N(CH2)2N(CH2COOH)2 powdery deposit on the crystals.
A compound that acts as a chelating effusion The ow of a gas through a
agent, reversibly binding with iron, mag- small aperture. The relative rates at which
nesium, and other metal ions. It is used in gases effuse, under the same conditions,
certain culture media bound with iron, is approximately inversely proportional to
which it slowly releases into the medium, the square roots of their densities.
and also in some forms of quantitative
analysis. egestion The expulsion from the body
of waste food materials that have never
EEG See electroencephalogram. left the gut, particularly the expulsion of
effective temperature See lumi- undigested materials from the gut
nosity. through the anus (see defecation). Eges-
tion should not be confused with *excre-
effective value See root-mean-square tion, in which the waste materials are
value.
produced by metabolic activity in the
effector A cell or organ that produces a bodys tissues.
physiological response when stimulated
egg 1. The fertilized ovum (*zygote) in
by a nerve impulse. Examples include
egg-laying animals, e.g. birds and insects,
muscles and glands.
after it emerges from the body. The egg is
effector neuron A nerve cell, such as a covered by *egg membranes that protect
motor neuron, that transmits impulses it from environmental damage, such as
egg membrane 268

drying. 2. (egg cell) The mature female Einstein coefcients Coefcients used
reproductive cell in animals and plants. in the *quantum theory of radiation, re-
See oosphere; ovum. lated to the probability of a transition oc-
curring between the ground state and an
egg membrane The layer of material
excited state (or vice versa) in the pro-
that covers an animal egg cell. Primary
cesses of *induced emission and *sponta-
membranes develop in the ovary and
neous emission. For an atom exposed to
cover the egg surface in addition to the
*electromagnetic radiation, the rate of ab-
normal plasma membrane. The primary
sorption Ra is given by Ra = B, where is
e membrane is called the vitelline mem-
brane in insects, molluscs, birds, and am-
the density of electromagnetic radiation
and B is the Einstein B coefcient associ-
phibians, the chorion in tunicates and
ated with absorption. The rate of induced
sh, and the zona pellucida in mammals.
emission is also given by B, with the
Insects have a second thicker membrane,
coefcient B of induced emission being
also called the chorion. Secondary mem-
equal to the coefcient of absorption. The
branes are secreted by the oviducts and
rate of spontaneous emission is given by
parts of the genital system while the egg
A, where A is the Einstein A coefcient of
is passing to the outside. They include the
spontaneous emission. The A and B coef-
jelly coat of frogs eggs and the albumen
cients are related by A = 8h3B/c3, where
and shell of birds eggs.
h is the *Planck constant, is the fre-
Ehrlich, Paul (18541915) German bac- quency of electromagnetic radiation, and
teriologist, who graduated as a physician c is the speed of light. The coefcients
in 1878. After working in a Berlin hospital were put forward by Albert Einstein in
for nine years he taught at Berlin Univer- 191617 in his analysis of the quantum
sity (unpaid because he was a Jew). In theory of radiation.
1890 he went to work with Robert Koch
Einstein equation 1. The massenergy
(18431910) to study tuberculosis, cholera,
relationship announced by Einstein in
and other diseases. In 1910 he discovered
1905 in the form E = mc2, where E is a
Salvarsan, an arsenical drug effective
quantity of energy, m its mass, and c is the
against syphilis. He was awarded the 1908
speed of light. It presents the concept that
Nobel Prize for physiology or medicine for
energy has mass. See also relativity.
his earlier work on serum therapy.
2. The relationship
eigenfunction An allowed *wave func-
Emax = hf W,
tion of a system in quantum mechanics.
The associated energies are eigenvalues. where Emax is the maximum kinetic en-
ergy of the electrons emitted in the photo-
Einstein, Albert (18791955) German-
emissive effect, h is the Planck constant, f
born US physicist, who took Swiss nation-
the frequency of the incident radiation,
ality in 1901. A year later he went to work
and W the *work function of the emitter.
in the Bern patent ofce. In 1905 he pub-
This is also written Emax = hf e, where e
lished ve enormously inuential papers,
is the electronic charge and a potential
one on *Brownian movement, one on the
difference, also called the work function.
*photoelectric effect, one on the special
(Sometimes W and are distinguished as
theory of *relativity, and one on energy
work function energy and work function
and inertia (which included the famous
potential.) The equation can also be ap-
expression E = mc2). In 1915 he published
plied to photoemission from gases, when
the general theory of relativity, concerned
it has the form: E = hf I, where I is the
mainly with gravitation. In 1921 he was
ionization potential of the gas.
awarded the Nobel Prize. In 1933, as a
Jew, Einstein decided to remain in the einsteinium Symbol Es. A radioactive
USA (where he was lecturing), as Hitler metallic transuranic element belonging to
had come to power. For the remainder of the *actinoids; a.n. 99; mass number of
his life he sought a unied eld theory. In the most stable isotope 254 (half-life 270
1939 he informed President Roosevelt days). Eleven isotopes are known. The el-
that an atom bomb was feasible and that ement was rst identied by Albert
Germany might be able to make one. Ghiorso and associates in debris from the
269 elastomer

rst hydrogen bomb explosion in 1952. after there is at rst a slight increase in
Microgram quantities of the element did strain with increased load until a point L
not become available until 1961. is reached. This is the elastic limit; up to
this point the deformation of the speci-
Einstein shift See redshift.
men is elastic, i.e. when the stress is re-
ejaculation The propulsion of semen moved the specimen returns to its
out of the erect penis due to powerful original length. Beyond the point L there
rhythmic contractions of the urethra. An is permanent deformation when the
ejaculation coincides with the peak of sex- stress is removed, i.e. the material has
ual excitement (orgasm) and is accompa- ceased to be elastic and has become plas- e
nied by various physiological effects in tic. In the plastic stages individual ma-
the body, such as increased respiration terials vary somewhat; in general,
rate and heart rate. however, at a point B there is a sudden in-
crease in strain with further increases of
ejecta Rocks and other material thrown stress this is the yield point. Beyond the
up when a crater is formed. Some craters point C, the breaking stress, the wire will
and their associated ejecta are volanic in snap (which occurs at point D).
origin, but most (especially on the moon,
Mercury, and other planets and their
satellites) are caused by the impact of me-
teorites. Often these are surrounded by an C
ejecta blanket, which is thickest near the elastic limit
breaking D
load

L B
craters rim. A stress
yield point breaking
Elasmobranchii See chondrichthyes. point
proportional limit
elastance The reciprocal of *capaci-
tance. It is measured in farad1 (some-
times called a daraf).
O strain
elastic cartilage See cartilage.
Elasticity
elastic collision A collision in which
the total kinetic energy of the colliding elastic modulus The ratio of the
bodies after collision is equal to their total *stress applied to a body to the *strain
kinetic energy before collision. Elastic col- produced. The Young modulus of elastic-
lisions occur only if there is no conversion ity, named after Thomas Young, refers to
of kinetic energy into other forms, as in longitudinal stress and strain. The bulk
the collision of atoms. In the case of modulus is the ratio of the pressure on a
macroscopic bodies this will not be the body to its fractional decrease in volume.
case as some of the energy will become The shear (or rigidity) modulus is the tan-
heat. In a collision between polyatomic gential force per unit area divided by the
molecules, some kinetic energy may angular deformation in radians.
be converted into vibrational and rota-
tional energy of the molecules, but other- elastin A brous protein that is the
wise molecular collisions appear to be major constituent of the yellow elastic
elastic. bres of *connective tissue. It is rich in
glycine, alanine, proline, and other non-
elastic bres See elastin. polar amino acids that are cross-linked,
elasticity The property of certain ma- making the protein relatively insoluble.
terials that enables them to return to Elastic bres can stretch to several times
their original dimensions after an applied their length and then return to their origi-
*stress has been removed. In general, if a nal size. Elastin is particularly abundant
stress is applied to a wire, the *strain will in elastic *cartilage, blood-vessel walls,
ligaments, and the heart.
increase in proportion (see OA on the illus-
tration) until a certain point called the elastomer A natural or synthetic
limit of proportionality is reached. This is rubber or rubberoid material, which
in accordance with *Hookes law. There- has the ability to undergo deformation
electret 270

under the inuence of a force and regain density) Symbol D. The charge per unit
its original shape once the force has been area that would be displaced across a
removed. layer of conductor placed across an *elec-
tric eld. This describes also the charge
electret A permanently electried sub-
density on an extended surface that could
stance or body that has opposite charges
be causing the eld.
at its extremities. Electrets resemble per-
manent magnets in many ways. An elec- electric eld A region in which an elec-
tret can be made by cooling certain waxes tric charge experiences a force usually be-
in a strong electric eld.
e cause of a distribution of other charges.
The electric eld strength or electric inten-
electrical energy A form of energy re-
sity (E) at any point in an electric eld is
lated to the position of an electric charge
dened as the force per unit charge ex-
in an electric eld. For a body with charge
perienced by a small charge placed at that
Q and an electric potential V, its electrical
point. This is equivalent to a potential gra-
energy is QV. If V is a potential difference,
dient along the eld and is measured in
the same expression gives the energy
volts per metre. The strength of the eld
transformed when the charge moves
can alternatively be described by its *elec-
through the p.d.
tric displacement D. The ratio D/E for
electric arc A luminous discharge be- measurements in a vacuum is the electric
tween two electrodes. The discharge constant 0. In a substance the observed
raises the electrodes to incandescence, the potential gradient is reduced by electron
resulting thermal ionization largely pro- movement so that D/E appears to increase:
viding the carriers to maintain the high the new ratio () is called the *permittiv-
current between the electrodes. ity of the substance. An electric eld can
electric-arc furnace A furnace used in be created by an isolated electric charge,
melting metals to make alloys, especially in which case the eld strength at a dis-
in steel manufacture, in which the heat tance r from a point charge Q is given by
source is an electric arc. In the direct-arc E = Q /4r2, where is the permittivity of
furnace, such as the Hroult furnace, an the intervening medium (see coulombs
arc is formed between the metal and an law). An electric eld can also be created
electrode. In the indirect-arc furnace, such by a changing magnetic eld.
as the Stassano furnace, the arc is formed
between two electrodes and the heat is ra-
diated onto the metal. bell

electric bell A device in which an elec-


hammer
tromagnetically operated hammer strikes
a bell (see illustration). Pressing the bell-
push closes a circuit, causing current to
ow from a battery or mains step-down light iron
trembler
transformer through an electromagnet.
The electromagnet attracts a piece of soft
iron attached to the hammer, which
strikes the bell and at the same time soft iron
core
breaks the circuit. The hammer springs
coil
back into its original position again, clos-
ing the circuit and causing the magnet to
attract the soft iron. This process contin- spring
ues until the bell-push is released. switch
electric charge See charge.
cell
electric constant See permittivity.
electric current See current.
electric displacement (electric ux Electric bell
271 electric potential

electric ux Symbol . In an *electric lights as less of the energy is converted


eld, the product of the electric ux den- into heat.
sity and the relevant area. See electric electric motor A machine for convert-
displacement. ing electrical energy into mechanical en-
electric ux density See electric dis- ergy. They are quiet, clean, and have a
placement. high efciency (7595%). They work on
the principle that a current passing
electricity Any effect resulting from the through a coil within a magnetic eld will
existence of stationary or moving electric
charges.
experience forces that can be used to ro-
tate the coil. In the induction motor, alter- e
electric lighting Illumination provided nating current is fed to a stationary coil
by electric currents. The devices used are (the stator), which both creates the mag-
the arc lamp, the light bulb (incandescent netic eld and induces a current in the ro-
lament lamp), and the uorescent tube. tating coil (rotor), which it surrounds. The
In the arc lamp, which is no longer used advantage of this kind of motor is that
as a general means of illumination, an current does not have to be fed through a
electric current ows through a gap be- commutator to a moving part. In the syn-
chronous motor, alternating current fed to
tween two carbon electrodes, between
the stator produces a magnetic eld that
which a high potential difference is main-
rotates and locks with the eld of the
tained. The current is carried by electrons
rotor, in this case an independent mag-
and ions in the vapour produced by the
net, causing the rotor to rotate at the
electrodes and a mechanism is required to
same speed as the stator eld rotates. The
bring the electrodes closer together as
rotor is either a permanent magnet or an
they are vaporized. The device produces a electromagnet fed by a direct current
strong white light but has many practical through slip rings. In the universal motor,
disadvantages. However, arcs enclosed in current is fed to the stator and, through a
an inert gas (usually xenon) are increas- commutator, to the rotor. In the series-
ingly used for such purposes as cinema wound motor the two are in series; in the
projectors. The common light bulb is a shunt-wound motor they are in parallel.
glass bulb containing a tungsten lament These motors can be used with either a.c.
and usually an inert gas. The passage of or d.c. but some small motors use a per-
an electric current through the lament manent magnet as the stator and require
heats it to a white heat. Inert gas is used d.c. for the rotor (via the commutator). See
in the bulb to minimize blackening of the also linear motor.
glass by evaporation of tungsten. In the
electric organ An organ occurring on
uorescent tube a glass tube containing
the body or tail of certain sh, such as the
mercury vapour (or some other gas) at a
electric ray (Torpedo) and electric eel (Elec-
low pressure has its inner surface coated
trophorus electricus). It gives an electric
with a uorescent substance. A discharge
shock when touched and is used either to
is created within the tube between two
stun prey or predators or, in some species,
electrodes. Electrons emitted by the cath- to maintain a weak electric eld in the
ode collide with gas atoms or molecules surrounding water that is used in naviga-
and raise them to an excited state (see ex- tion. The organ is composed of modied
citation). When they fall back to the muscle cells (electroplate cells), nervous
*ground state they emit photons of ultra- stimulation of which greatly increases the
violet radiation, which is converted to vis- potential difference across the cell. The
ible light by the coating of phosphor on electroplates are in series so a high overall
the inner walls of the tube. In some voltage can be achieved.
lamps, such as the *sodium-vapour and
*mercury-vapour lamps used in street electric polarization See dielectric.
lighting, no uorescent substance is used, electric potential Symbol V. The en-
the light being emitted directly by the ex- ergy required to bring unit electric charge
cited atoms of sodium or mercury. Vapour from innity to the point in an electric
lights are more efcient than lament eld at which the potential is being
electric power 272

specied. The unit of electric potential is cathode is the negative electrode. 2. See
the volt. The potential difference (p.d.) be- half cell.
tween two points in an electric eld or cir-
electrodeposition The process of
cuit is the difference in the values of the
depositing one metal on another by elec-
electric potentials at the two points, i.e. it
trolysis, as in *electroforming and
is the work done in moving unit charge
*electroplating.
from one point to the other.
electrode potential The potential dif-
electric power The rate of expending
ference produced between the electrode
e energy or doing work in an electrical sys-
tem. For a direct-current circuit, it is given
and the solution in a *half cell. It is not
possible to measure this directly since any
by the product of the current passing
measurement involves completing the cir-
through a system and the potential differ-
cuit with the electrolyte, thereby intro-
ence across it. In alternating-current cir-
cuits, the power is given by VIcos, where ducing another half cell. Standard
electrode potentials E are dened by
V and I are the RMS values and is the
*phase angle. Cos is called the power measuring the potential relative to a stan-
factor of the circuit. dard *hydrogen half cell using 1.0 molar
solution at 25C. The convention is to des-
electric spark The transient passage of ignate the cell so that the oxidized form is
an electric current through a gas between written rst. For example,
two points of high opposite potential,
with the emission of light and sound. Pt(s)|H2(g)H+(aq)|Zn2+(aq)|Zn(s)
*Lightning consists of a spark between a The e.m.f. of this cell is 0.76 volt (i.e. the
cloud and earth or between two oppo- zinc electrode is negative). Thus the stan-
sitely charged parts of the same cloud. dard electrode potential of the Zn2+|Zn
electric susceptibility See susceptibil- half cell is 0.76 V. Electrode potentials
ity. are also called reduction potentials. See
also electromotive series.
electrocardiogram (ECG) A tracing or
graph of the electrical activity of the electrodialysis A method of obtaining
heart. Recordings are made from elec- pure water from water containing a salt,
trodes fastened over the heart and usually as in *desalination. The water to be
on both arms and a leg. Changes in the puried is fed into a cell containing two
normal pattern of an ECG may indicate electrodes. Between the electrodes is
heart irregularities or disease. placed an array of *semipermeable mem-
branes alternately semipermeable to posi-
electrochemical cell See cell. tive ions and negative ions. The ions tend
electrochemical equivalent Symbol to segregate between alternate pairs of
z. The mass of a given element liberated membranes, leaving pure water in the
from a solution of its ions in electrolysis other gaps between membranes. In this
by one coulomb of charge. See faradays way, the feed water is separated into two
laws (of electrolysis). streams: one of pure water and the other
of more concentrated solution.
electrochemical series See electro-
motive series. electrodynamics The study of electric
charges in motion, the forces created by
electrochemistry The study of chemi- electric and magnetic elds, and the rela-
cal properties and reactions involving ions tionship between them. Compare electro-
in solution, including electrolysis and statics.
electric cells.
electroencephalogram (EEG) A trac-
electrochromatography See electro-
ing or graph of the electrical activity of
phoresis.
the brain. Electrodes taped to the scalp
electrode 1. A conductor that emits or record electrical waves from different
collects electrons in a cell, thermionic parts of the brain. The pattern of an EEG
valve, semiconductor device, etc. The reects an individuals level of conscious-
anode is the positive electrode and the ness and can be used to detect such disor-
273 electromagnetic induction

ders as epilepsy, tumours, or brain dam- cell can be made to pass current easily
age. See also brain death. in one direction but hardly at all in the
other. Examples include a lead
electroforming A method of forming
intricate metal articles or parts by *elec- aluminium cell with ammonium phos-
trodeposition of the metal on a removable phate(V) electrolyte and a tantalumlead
conductive mould. cell with sulphuric acid as the electrolyte.

electroluminescence See lumines- electrolytic rening The purication


cence. of metals by electrolysis. It is commonly

electrolysis The production of a chemi-


applied to copper. A large piece of impure
copper is used as the anode with a thin
e
cal reaction by passing an electric current strip of pure copper as the cathode. Cop-
through an electrolyte. In electrolysis, per(II) sulphate solution is the electrolyte.
positive ions migrate to the cathode and Copper dissolves at the anode: Cu Cu2+
negative ions to the anode. The reactions + 2e, and is deposited at the cathode. The
occurring depend on electron transfer at
net result is transfer of pure copper from
the electrodes and are therefore redox re-
anode to cathode. Gold and silver in the
actions. At the anode, negative ions in so-
impure copper form a so-called anode
lution may lose electrons to form neutral
sludge at the bottom of the cell, which is
species. Alternatively, atoms of the elec-
recovered.
trode can lose electrons and go into solu-
tion as positive ions. In either case the electrolytic separation A method of
reaction is an oxidation. At the cathode, separating isotopes by exploiting the dif-
positive ions in solution can gain elec- ferent rates at which they are released in
trons to form neutral species. Thus cath- electrolysis. It was formerly used for sepa-
ode reactions are reductions. rating deuterium and hydrogen. On elec-
electrolyte A liquid that conducts elec- trolysis of water, hydrogen is formed at
tricity as a result of the presence of posi- the cathode more readily than deuterium,
tive or negative ions. Electrolytes are thus the water becomes enriched with
molten ionic compounds or solutions con- deuterium oxide.
taining ions, i.e. solutions of ionic salts or electromagnet A magnet consisting of
of compounds that ionize in solution. Liq- a soft ferromagnetic core with a coil of in-
uid metals, in which the conduction is by sulated wire wound round it. When a cur-
free electrons, are not usually regarded as rent ows through the wire the core
electrolytes. Solid conductors of ions, as in becomes magnetized; when the current
the sodiumsulphur cell, are also known
ceases to ow the core loses its magnetiza-
as electrolytes.
tion. Electromagnets are used in switches,
electrolytic capacitor See capacitor. solenoids, electric bells, metal-lifting
cranes, and many other applications.
electrolytic cell A cell in which elec-
trolysis occurs; i.e. one in which current is electromagnetic induction The pro-
passed through the electrolyte from an duction of an electromotive force in a
external source. conductor when there is a change of mag-
electrolytic corrosion Corrosion that netic ux linkage with the conductor or
occurs through an electrochemical reac- when there is relative motion of the con-
tion. See rusting. ductor across a magnetic eld. The magni-
tude of the e.m.f. is proportional (and in
electrolytic gas (detonating gas) The modern systems of units equal) to the rate
highly explosive gas formed by the elec- of change of the ux linkage or the rate of
trolysis of water. It consists of two parts cutting ux d/dt; the sense of the in-
hydrogen and one part oxygen by volume. duced e.m.f. is such that any induced cur-
electrolytic rectier A *rectier con- rent opposes the change causing the
sisting of two dissimilar electrodes im- induction, i.e. E = d/dt. See faradays
mersed in an electrolyte. By suitable laws; lenzs law; neumanns law; in-
choice of electrodes and electrolyte the ductance.
electromagnetic interaction 274

electromagnetic interaction See fun- (1 centimetre in diameter), exerts a force


damental interactions. of 1 dyne on unit magnetic pole at the
centre of the circle. In e.m.u. the mag-
electromagnetic pump A pump used
netic constant is of unit magnitude. The
for moving liquid metals, such as the liq-
system has now been replaced by *SI
uid-sodium coolant in a fast nuclear reac-
units for most purposes. Compare electro-
tor. The liquid is passed through a
static units; gaussian units; heaviside
attened pipe over two electrodes be-
lorentz units.
tween which a direct current ows. A
magnetic eld at right angles to the cur-
e rent causes a force to be created directly
electromagnetic wave See electro-
magnetic radiation; wave.
on the liquid, along the axis of the tube.
electrometallurgy The uses of electri-
The pump has no moving parts and is
cal processes in the separation of metals
therefore safe and trouble free.
from their ores, the rening of metals, or
electromagnetic radiation Energy re- the forming or plating of metals.
sulting from the acceleration of electric
charge and the associated electric elds electrometer A measuring instrument
and magnetic elds. The energy can be re- for determining a voltage difference with-
garded as waves propagated through out drawing an appreciable current from
space (requiring no supporting medium) the source. Originally electrostatic instru-
involving oscillating electric and magnetic ments based on the electroscope, they are
elds at right angles to each other and to now usually based on operational am-
the direction of propagation. In a vacuum pliers, solid-state devices with high input
the waves travel with a constant speed impedances. Electrometers are also used
(the speed of light) of 2.9979 108 metres to measure low currents (nanoamperes),
per second; if material is present they are by passing the current through a high re-
slower. Alternatively, the energy can be sistance.
regarded as a stream of *photons travel- electromotive force (e.m.f.) The great-
ling at the speed of light, each photon est potential difference that can be gener-
having an energy hc/, where h is the ated by a particular source of electric
Planck constant, c is the speed of light, current. In practice this may be observ-
and is the wavelength of the associated able only when the source is not supply-
wave. A fusion of these apparently ing current, because of its *internal
conicting concepts is possible using the resistance.
methods of *quantum mechanics or
*wave mechanics. The characteristics of electromotive series (electrochemical
the radiation depend on its wavelength. series) A series of chemical elements ar-
See electromagnetic spectrum. ranged in order of their *electrode poten-
tials. The hydrogen electrode (H+ + e
electromagnetic spectrum The range H2) is taken as having zero electrode po-
of wavelengths over which *electromag- tential. Elements that have a greater ten-
netic radiation extends. The longest waves dency than hydrogen to lose electrons to
(105103 metres) are radio waves, the their solution are taken as electropositive;
next longest (103106 m) are infrared those that gain electrons from their solu-
waves, then comes the narrow band tion are below hydrogen in the series and
(47 107 m) of visible light, followed by are called electronegative. The series
ultraviolet waves (107109 m), X-rays shows the order in which metals replace
(1091011 m), and gamma rays (1011 one another from their salts; electroposi-
1014 m). tive metals will replace hydrogen from
electromagnetic units (e.m.u.) A sys- acids. The chief metals and hydrogen,
tem of electrical units formerly used in placed in order in the series, are: potas-
the *c.g.s. system. The e.m.u. of electric sium, calcium, sodium, magnesium, alu-
current is the abampere (all e.m.u. have minium, zinc, cadmium, iron, nickel, tin,
the prex ab- attached to the names of lead, hydrogen, copper, mercury, silver,
practical units). The abampere is the cur- platinum, gold. This type of series is
rent that, owing in an arc of a circle sometimes referred to as an activity series.
275 electronegative

electron An *elementary particle, tial difference V acquires a kinetic energy


classed as a *lepton, with a rest mass mv2/2 = eV, where v is the velocity of the
(symbol me) of 9.109 3897(54) 1031 kg electron. The (nonrelativistic) momentum
and a negative charge of 1.602 177 33(49) (p) of the electron is (2eVm). The de
1019 coulomb. Electrons are present in Broglie wavelength () of an electron is
all atoms in groupings called shells given by h/p, where h is the Planck con-
around the nucleus; when they are de- stant, thus = h/(2eVm). For an accelerat-
tached from the atom they are called free ing voltage of 3600 V, the wavelength of
electrons. The antiparticle of the electron the electron beam is 0.02 nanometre,
is the positron. some 3 104 times shorter than visible ra- e
diation.
electron afnity Symbol A. The energy
Electrons then, like X-rays, show
change occurring when an atom or mol-
diffraction effects with molecules and
ecule gains an electron to form a negative
crystals in which the interatomic spacing
ion. For an atom or molecule X, it is
is comparable to the wavelength of the
the energy released for the electron-
beam. They have the advantage that their
attachment reaction
wavelength can be set by adjusting the
X(g) + e X(g) voltage. Unlike X-rays they have very low
penetrating power. The rst observation
Often this is measured in electronvolts.
of electron diffraction was by George
Alternatively, the molar enthalpy change,
Thomson (18921975) in 1927, in an ex-
H, can be used.
periment in which he passed a beam of
electron biprism An arrangement of electrons in a vacuum through a very thin
elds that splits a beam of electrons or gold foil onto a photographic plate. Con-
other charged particles in an analogous centric circles were produced by diffrac-
way to an optical biprism. tion of electrons by the lattice. The same
year Clinton J. Davisson (18811958) and
electron capture 1. The formation of a
Lester Germer (18961971) performed a
negative ion by an atom or molecule
classic experiment in which they obtained
when it acquires an extra free electron.
diffraction patterns by glancing an elec-
2. A radioactive transformation in which
tron beam off the surface of a nickel crys-
a nucleus acquires an electron from an
tal. Both experiments were important
inner orbit of the atom, thereby trans-
verications of de Broglies theory and the
forming, initially, into a nucleus with the
new quantum theory.
same mass number but an atomic number
Electron diffraction, because of the low
one less than that of the original nucleus
penetration, cannot easily be used to in-
(capture of the electron transforms a pro-
vestigate crystal structure. It is, however,
ton into a neutron). This type of capture is
employed to measure bond lengths and
accompanied by emission of an X-ray pho-
angles of molecules in gases. Moreover, it
ton or Auger electron as the vacancy in
is extensively used in the study of solid
the inner orbit is lled by an outer elec-
surfaces and absorption. The main tech-
tron.
niques are low-energy electron diffraction
electron-decient compound A com- (LEED) in which the electron beam is
pound in which there are fewer electrons reected onto a uorescent screen, and
forming the chemical bonds than re- high-energy electron diffraction (HEED)
quired in normal electron-pair bonds. used either with reection or transmis-
Such compounds use *multicentre bonds. sion in investigating thin lms.
See borane.
electronegative Describing elements
electron diffraction *Diffraction of a that tend to gain electrons and form nega-
beam of electrons by atoms or molecules. tive ions. The halogens are typical elec-
The fact that electrons can be diffracted tronegative elements. For example, in
in a similar way to light and X-rays shows hydrogen chloride, the chlorine atom is
that particles can act as waves (see de more electronegative than the hydrogen
broglie wavelength). An electron (mass and the molecule is polar, with negative
m, charge e) accelerated through a poten- charge on the chlorine atom. There are
electron flow 276

various ways of assigning values for the coils to produce a magnetic eld or an ar-
electronegativity of an element. Mulliken rangement of electrodes between which
electronegativities are calculated from E = an electric eld is created. Electron lenses
(I + A)/2, where I is ionization potential are used in *electron microscopes and
and A is electron afnity. More com- *cathode-ray tubes.
monly, Pauling electronegativities are electron microscope A form of micro-
used. These are based on bond dissocia- scope that uses a beam of electrons in-
tion energies using a scale in which stead of a beam of light (as in the optical
uorine, the most electronegative el-
e ement, has a value 4. Some other values
microscope) to form a large image of a
very small object. In optical microscopes
on this scale are B 2, C 2.5, N 3.0, O 3.5, the resolution is limited by the wave-
Si 1.8, P 2.1, S 2.5, Cl 3.0, Br 2.8. length of the light. High-energy electrons,
electron ow The transfer of electrons however, can be associated with a consid-
along a series of carrier molecules in the erably shorter wavelength than light; for
*electron transport chain. example, electrons accelerated to an en-
ergy of 105 electronvolts have a wave-
electron gun A device used in *cath- length of 0.004 nanometre (see de broglie
ode-ray tubes (including television tubes), wavelength) enabling a resolution of
electron microscopes, etc., to produce a 0.20.5 nm to be achieved. The transmis-
steady narrow beam of electrons. It usu- sion electron microscope (see illustration)
ally consists of a heated cathode, control has an electron beam, sharply focused by
grid, and two or more annular anodes in- *electron lenses, passing through a very
serted in an evacuated tube. The electrons thin metallized specimen (less than 50
emitted by the cathode are attracted to nanometres thick) onto a uorescent
the nal anode, through which they pass. screen, where a visual image is formed.
The intensity of the beam is regulated by This image can be photographed. The
the control grid and potential differences scanning electron microscope can be used
between the anodes create electric elds with thicker specimens and forms a per-
that focus the diverging electrons into a spective image, although the resolution
narrow beam. and magnication are lower. In this type
electronic mail (e-mail) Messages, docu-
ments, etc., sent between users of com-
puter systems, the computer systems
being used to transport and hold the source of electrons
e-mail. The service itself is also referred to
as electronic mail. The sender and recipi-
ent(s) need not be at their computers at
the same time to communicate, and the condenser lens
computer systems may be situated world- object
wide. The sender creates an e-mail by
means of a mail-sending computer pro- objective lens
gram, and a mail transport system then
takes responsibility for delivering the
e-mail to the indicated address(es).
electronics The study and design of
intermediate image
control, communication, and computing
devices that rely on the movement of
electrons in circuits containing semicon- projector lens
ductors, thermionic valves, resistors, ca-
pacitors, and inductors. See Chronology.
electron lens A device used to focus an
electron beam. It is analogous to an opti- final image
cal lens but instead of using a refracting
material, such as glass, it uses a coil or Principle of transmission electron microscope
277

ELECTRONICS

1887 Radio waves are discovered by Heinrich Hertz.


1894 Oliver Lodge invents the coherer for detecting radio waves.
Marconi develops radio telegraphy.
1897 J. J. Thomson discovers the electron.
1902
1903
US engineer Reginald Fessenden (18661932) develops radio telephony.
Danish engineer Valdemar Poulsen (18691942) invents the arc transmitter
e
for radio telegraphy.
1904 British engineer Ambrose Fleming (18491945) invents the diode
thermionic valve.
1906 US engineer Lee De Forest (18731961) invents the triode thermionic valve.
US electrical engineer Greenleaf Pickard (18771956) patents the crystal
detector for radios.
Fessenden introduces amplitude modulation in radio broadcasting.
1911 German physicist Karl Braun (18501918) invents cathode-ray tube
scanning.
1912 Fessenden develops the heterodyne radio receiver.
1919 US electrical engineer Edwin Armstrong (18901954) develops the
superheterodyne radio receiver.
1921 US physicist Albert Hull (18801966) invents the magnetron microwave-
generating valve.
1923 Russian-born US engineer Vladimir Zworykin (18891982) invents the
iconoscope television camera-tube.
1928 Scottish inventor John Logie Baird (18881946) and Vladimir Zworykin
independently develop television.
1930 Swedish-born US electronics engineer Ernst Alexanderson (18781975)
invents an all-electronic television system.
1933 US electrical engineer Edwin Armstrong (18901954) develops frequency
modulation radio broadcasting.
1947 US physicists John Bardeen, Walter Brattain (190287), and William
Shockley (191089) invent the point-contact transistor.
1950 US engineers develop the Videcon television camera tube.
1953 Chinese-born US computer engineer An Wang (192090) invents the
magnetic core computer memory.
1954 US physicist Charles Townes (1915 ) and Soviet physicists Nikolai Basov
and Aleksandr Prokhorov (1916 ) independently develop the maser.
1958 US electronics engineers Jack Kilby and Robert Noyce (192790) develop
integrated circuits.
1960 US physicist Theodore Maiman (1927 ) invents the ruby laser.
1961 US electronics engineer Steven Hofstein develops the field-effect
transistor.
1971 US electronics engineer Marcian Edward Hoff (1937 ) designs the first
microprocessor (Intel 4004).
1977 US engineers transmit television signals along optical fibres.
electron optics 278

of instrument a beam of primary elec- Electron-transfer reactions often involve


trons scans the specimen and those that complexes of transition metals. In such
are reected, together with any secondary complexes one general mechanism for
electrons emitted, are collected. This cur- electron transfer is the inner-sphere mech-
rent is used to modulate a separate elec- anism, in which two complexes form an
tron beam in a TV monitor, which scans intermediate, with ligand bridges en-
the screen at the same frequency, conse- abling electrons to be transferred from
quently building up a picture of the speci- one complex to another complex. The
men. The resolution is limited to about other main mechanism is the outer-
e 1020 nm. See also field-emission micro- sphere mechanism, in which two com-
scope; field-ionization microscope. plexes retain all their ligands, with
electrons passing from one complex to
electron optics The study of the use of
the other. The rates of electron-transfer
*electron lenses in the *electron micro-
reactions vary enormously. These rates
scope, *cathode-ray tubes, and other simi-
can be explained in terms of the way in
lar devices. The focusing of beams of
which molecules of the solvent solvating
positive or negative ions also relies on
the reactants rearrange so as to solvate
these methods.
the products in the case of the outer-
electron probe microanalysis (EPM) sphere mechanism. In the case of the
A method of analysing a very small quan- inner-sphere (ligand-bridged) reactions the
tity of a substance (as little as 1013 gram). rate of the reaction depends on the inter-
The method consists of directing a very mediate and the way in which the elec-
nely focused beam of electrons on to the tron is transferred.
sample to produce the characteristic X-ray
electron transport chain (electron
spectrum of the elements present. It can
transport system) A sequence of biochem-
be used quantitatively for elements with
ical reductionoxidation reactions that
atomic numbers in excess of 11.
effects the transfer of electrons through a
electron-spin resonance (ESR) A spec- series of carriers. An electron transport
troscopic method of locating electrons chain, also known as the respiratory
within the molecules of a paramagnetic chain, forms the nal stage of *aerobic
substance (see magnetism) in order to pro- respiration. It results in the transfer of
vide information regarding its bonds and electrons or hydrogen atoms derived from
structure. The spin of an unpaired elec- the *Krebs cycle to molecular oxygen,
tron is associated with a *magnetic mo- with the formation of water. At the same
ment that is able to align itself in one of time it conserves energy from food or
two ways with an applied external mag- light in the form of *ATP. The chain com-
netic eld. These two alignments corre- prises a series of *carrier molecules that
spond to different *energy levels, with a undergo reversible reductionoxidation
statistical probability, at normal tempera- reactions, accepting electrons and then
tures, that there will be slightly more in donating them to the next carrier in the
the lower state than in the higher. By ap- chain a process known as electron ow.
plying microwave radiation to the sample In the mitochondria, NADH and FADH2,
a transition to the higher state can be generated by the Krebs cycle, transfer
achieved. The precise energy difference their electrons to a chain comprising
between the two states of an electron de- avin mononucleotide (FMN), *ubi-
pends on the surrounding electrons in the quinone, and a series of *cytochromes.
atom or molecule. In this way the position This process is coupled to the formation
of unpaired electrons can be investigated. of ATP at three sites along the chain (see
The technique is used particularly in oxidative phosphorylation). The ATP is
studying free radicals and paramagnetic then carried across the mitochondrial
substances such as inorganic complexes. membrane in exchange for ADP. An
See also nuclear magnetic resonance. electron transport chain also occurs in
*photosynthesis.
electron-transfer reaction A chemi-
cal reaction that involves the transfer, ad- electronvolt Symbol eV. A unit of en-
dition, or removal of electrons. ergy equal to the work done on an elec-
279 electrostatic generator

tron in moving it through a potential dif- paper, soaked in a buffer with two elec-
ference of one volt. It is used as a measure trodes making contact. The sample is
of particle energies although it is not an placed between the electrodes and a volt-
*SI unit. 1 eV = 1.602 1019 joule. age applied. Different components of the
mixture migrate at different rates, so the
electroorganic reaction An organic
sample separates into zones. The compo-
reaction produced in an electrolytic cell.
nents can be identied by the rate at
Electroorganic reactions are used to syn-
which they move. In gel electrophoresis
thesize compounds that are difcult to
the medium is a gel, typically made of
produce by conventional techniques. An
example of an electroorganic reaction is
polyacrylamide, agarose, or starch. e
Electrophoresis, which has also been
*Kolbes method of synthesizing alkanes.
called electrochromatography, is used ex-
electrophile An ion or molecule that is tensively in studying mixtures of proteins,
electron decient and can accept elec- nucleic acids, carbohydrates, enzymes,
trons. Electrophiles are often reducing etc. In clinical medicine it is used for de-
agents and Lewis *acids. They are either termining the protein content of body
positive ions (e.g. NO2+) or molecules that uids.
have a positive charge on a particular electrophorus An early form of *elec-
atom (e.g. SO3, which has an electron- trostatic generator. It consists of a at di-
decient sulphur atom). In organic reac- electric plate and a metal plate with an
tions they tend to attack negatively insulated handle. The dielectric plate is
charged parts of a molecule. Compare nu- charged by friction and the metal plate is
cleophile. placed on it and momentarily earthed,
electrophilic addition An *addition which leaves the metal plate with an in-
reaction in which the rst step is attack duced charge of opposite polarity to that
by an electrophile (e.g. a positive ion) on of the dielectric plate. The process can be
an electron-rich part of the molecule. An repeated until all of the original charge
example is addition to the double bonds has leaked away.
in alkenes. electroplating A method of plating one
electrophilic substitution A *substi- metal with another by *electrodeposition.
tution reaction in which the rst step is The articles to be plated are made the cath-
attack by an electrophile. Electrophilic ode of an electrolytic cell and a rod or bar
substitution is a feature of reactions of of the plating metal is made the anode.
benzene (and its compounds) in which a Electroplating is used for covering metal
positive ion approaches the delocalized pi with a decorative, more expensive, or cor-
electrons on the benzene ring. rosion-resistant layer of another metal.

electrophoresis (cataphoresis) A tech- electropositive Describing elements


nique for the analysis and separation of that tend to lose electrons and form posi-
colloids, based on the movement of tive ions. The alkali metals are typical
charged colloidal particles in an electric electropositive elements.
eld. There are various experimental electroscope A device for detecting
methods. In one the sample is placed in a electric charge and for identifying its po-
U-tube and a buffer solution added to larity. In the gold-leaf electroscope two
each arm, so that there are sharp bound- rectangular gold leaves are attached to the
aries between buffer and sample. An elec- end of a conducting rod held in an insu-
trode is placed in each arm, a voltage lated frame. When a charge is applied to a
applied, and the motion of the boundaries plate attached to the other end of the con-
under the inuence of the eld is ob- ducting rod, the leaves move apart owing
served. The rate of migration of the parti- to the mutual repulsion of the like
cles depends on the eld, the charge on charges they have received.
the particles, and on other factors, such as
electrostatic eld The *electric eld
the size and shape of the particles. More
that surrounds a stationary charged body.
simply, electrophoresis can be carried out
using an adsorbent, such as a strip of lter electrostatic generator A device used
electrostatic precipitation 280

to build up electric charge to an extreme bergSalam model or WS model. Because


potential usually for experimental pur- early developments of these ideas were
poses. The *electrophorus and the put forward by Sheldon Glashow, it is
*Wimshurst machine were early exam- sometimes known as the Glashow
ples; a more usual device now is the *Van WeinbergSalam model or GWS model. In
de Graaff generator. this electroweak theory the gauge group
is non-Abelian and the gauge symmetry is
electrostatic precipitation A method
a *broken symmetry. The electroweak in-
of removing solid and liquid particles
teraction is mediated by photons and by
e from suspension in a gas. The gas is ex-
posed to an electric eld so that the parti- intermediate vector bosons, called the
cles are attracted to and deposited on a *W boson and the *Z boson. The observa-
suitably placed electrode. Electrostatic tion of these particles in 1983/84, with
precipitation is widely used to remove their predicted energies, was a major suc-
dust and other pollutants from waste cess of the theory. The theory successfully
gases and from air. See also cottrell pre- accounts for existing data for electroweak
cipitator. processes and also predicts the existence
of a heavy particle with spin 0, the *Higgs
electrostatics The study of electric boson.
charges at rest, the forces between them
(see coulombs law), and the electric elds electrum 1. An alloy of gold and silver
associated with them. Compare electrody- containing 5588% of gold. 2. A *German
namics. silver alloy containing 52% copper, 26%
nickel, and 22% zinc.
electrostatic units (e.s.u.) A system of
electrical units in the *c.g.s. system. The element A substance that cannot be de-
e.s.u. of electric charge is the statcoulomb composed into simpler substances. In an
(all e.s.u. have the prex stat- attached to element, all the atoms have the same
the names of practical units). The stat- number of protons or electrons, although
coulomb is the quantity of electric charge the number of neutrons may vary. There
that will repel an equal quantity 1 centi- are 92 naturally occurring elements. See
metre distant with a force of 1 dyne. In also periodic table; transuranic el-
e.s.u. the electric constant is of unit mag- ements; transactinide elements.
nitude. The system has now been replaced elementary particles The fundamen-
for most purposes by *SI units. Compare tal constituents of all the matter in the
electromagnetic units; gaussian units; universe. By the beginning of the 20th
heavisidelorentz units. century, the electron and the proton had
electrostriction A change in the di- been discovered, but it was not until 1932
mensions of a body as a result of reorien- that the existence of the neutron was
tation of its molecules when it is placed in denitely established. Since 1932, it had
an electric eld. If the eld is not homo- been known that atomic nuclei consist of
geneous the body will tend to move; if its both protons and neutrons (except hydro-
relative permittivity is higher than that of gen, whose nucleus consists of a lone pro-
its surroundings it will tend to move into ton). Between 1900 and 1930, *quantum
a region of higher eld strength. Compare mechanics was also making progress in
magnetostriction. the understanding of physics on the
atomic scale. Non-relativistic quantum
electrovalent bond See chemical theory was completed in an astonishingly
bond. brief period (192326), but it was the rela-
electroweak theory A *gauge theory tivistic version that made the greatest im-
(sometimes called quantum avour- pact on our understanding of elementary
dynamics, or QFD) that gives a unied de- particles. Diracs discovery in 1928 of the
scription of the electromagnetic and weak equation that bears his name led to the
interactions (see fundamental interac- discovery of the positive electron or
tions). A successful electroweak theory *positron. The mass of the positron is
was proposed in 1967 by Steven Weinberg equal to that of the negative electron
and Abdus Salam, known as the Wein- while its charge is equal in magnitude but
281 elementary particles

opposite in sign. Pairs of particles related mass was discovered in *cosmic radiation.
to each other in this way are said to be This particle was named the -meson or
antiparticles of each other. Positrons have muon. The has a charge equal to the
only a transitory existence; that is, they electron, and its antiparticle + has a posi-
do not form part of ordinary matter. tive charge of equal magnitude. However,
Positrons and electrons are produced si- physicists soon discovered that muons do
multaneously in high-energy collisions of not interact with nuclear particles suf-
charged particles or gamma rays with ciently strongly to be Yukawas meson. It
matter in a process called *pair produc- was not until 1947 that a family of me-
tion. sons with the appropriate properties was e
The union of *relativity and quantum discovered. These were the -mesons or
mechanics therefore led to speculation as pions, which occur in three types: posi-
early as 1932 that there might also be an- tive, negative, and neutral. Pions, which
tiprotons and antineutrons, bearing a sim- interact strongly with nuclei, have in fact
ilar relationship to their respective turned out to be the particles predicted by
ordinary particles as the positron does to Yukawa in the 1930s. The nuclear force
the electron. However, it was not until between protons and neutrons was given
1955 that particle beams were made suf- the name strong interaction (see funda-
ciently energetic to enable these antimat- mental interactions) and until the
ter particles to be observed. It is now 1960s it was thought to be an exchange
understood that all known particles have force as proposed by Yukawa.
antimatter equivalents, which are pre- A theory of the weak interaction was
dicted by relativistic quantum equations. also in its infancy in the 1930s. The weak
By the mid-1930s the list of known and interaction is responsible for *beta decay,
theoretically postulated particles was still in which a radioactive nucleus is trans-
small but steadily growing. At this time formed into a slightly lighter nucleus
the Japanese physicist Hideki Yukawa with the emission of an electron. How-
(190781) was studying the possible *fun- ever, beta decays posed a problem be-
damental interactions that could hold the cause they appeared not to conserve
nucleus together. Since the nucleus is a energy and momentum. In 1931 *Pauli
closely packed collection of positively proposed the existence of a neutral parti-
charged protons and neutral neutrons, cle that might be able to carry off the
clearly it could not be held together by an missing energy and momentum in a beta
electromagnetic force; there had to be a decay and escape undetected. Three years
different and very large force capable of later, *Fermi included Paulis particle in a
holding proton charges together at such comprehensive theory of beta decay,
close proximity. This force would neces- which seemed to explain many experi-
sarily be restricted to the short range of mentally observed results. Fermi called
nuclear dimensions, because evidence of this new particle the *neutrino, the exis-
its existence only arose after the discovery tence of which was nally established in
of the constituents of the atomic nucleus. the 1950s.
Guided by the properties required of this A plethora of experiments involving the
new force, Yukawa proposed the exis- neutrino revealed some remarkable prop-
tence of a particle called the *meson, erties for this new particle. The neutrino
which was responsible for transmitting was found to have an intimate connection
nuclear forces. He suggested that protons with the electron and muon, and indeed
and neutrons in the nucleus could inter- never appeared without the simultaneous
act by emitting and absorbing mesons. For appearance of one or other of these parti-
this reason this new type of force was cles. A conservation law was postulated to
called an *exchange force. Yukawa was explain this observation. Numbers were
even able to predict the mass of his assigned to the electron, muon, and neu-
meson (meaning middle weight), which trino, so that during interactions these
turned out to be intermediate between numbers were conserved; i.e. their alge-
the proton and the electron. braic sums before and after these interac-
Only a year after Yukawa had made this tions were equal. Since these particles
suggestion, a particle of intermediate were among the lightest known at the
elements of an orbit 282

time, these assigned numbers became masses, which is indicated by the use of
known as lepton numbers (lepton: light asterisks in the table below.
ones). In order to make the assignments In the 1960s, the development of high-
of lepton number agree with experiment, energy accelerators and more sophisti-
it is necessary to postulate the existence cated detection systems led to the
of two types of neutrino. Each of these discovery of many new and exotic parti-
types is associated with either the elec- cles. They were all unstable and existed
tron or muon; there are thus muon neu- for only small fractions of a second; never-
trinos and electron neutrinos. In 1978 the theless they set into motion a search for a
e tau particle or tauon was discovered and theoretical description that could account
was added to the list of particles with as- for them all. The large number of these
signed lepton numbers. The conservation apparently fundamental particles sug-
of lepton number in the various interac- gested strongly that they do not, in fact,
tions involving the tau requires the exis- represent the most fundamental level of
tence of an equivalent tau neutrino. The the structure of matter. Physicists found
six particles with assigned lepton num- themselves in a position similar to
bers are now known as *leptons. Mendeleev when the *periodic table was
Neutrinos have zero charge and were being developed. Mendeleev realized that
originally thought to have zero rest mass,
there had to be a level of structure below
but there has been increasing indirect ex-
the elements themselves, which ex-
perimental evidence to the contrary. In
plained the chemical properties and the
1985 a Soviet team reported a measure-
interrelations between elements.
ment, for the rst time, of a non-zero neu-
Murray Gell-Mann and his collaborators
trino mass. The mass measured was
proposed the particle-physics equivalent
extremely small (10 000 times less than
of the periodic table in 1961. In this struc-
the mass of the electron), but subsequent
attempts independently to reproduce ture, leptons were indeed regarded as fun-
these results did not succeed. More re- damental particles, but the short-lived
cently (199899), Japanese and US groups particles discovered in the 1960s were
have put forward theories and corroborat- not. These particles were found to
ing experimental evidence to suggest, in- undergo strong interactions, which did
directly, that neutrinos do have mass. In not seem to affect the leptons. Gell-Mann
these experiments neutrinos are found to called these strongly interacting particles
apparently disappear. Since it is unlikely the *hadrons and proposed that they oc-
that momentum and energy are actually curred in two different types: baryons and
vanishing from the universe, a more plau- mesons. These two different types cor-
sible explanation is that the types of neu- responded to the two different ways of
trinos detected are changing into types constructing hadrons from constituent
that cannot be detected. Present theoreti- particles, which Gell-Mann called quarks.
cal considerations imply that the masses These quarks came in three avours, up
of neutrinos involved cannot be equal to (u), down (d), and strange (s). These three
one another, and therefore they cannot quarks were thought to be the fundamen-
all be zero. This speculative work has not tal constituents of hadrons, i.e. matter
yet yielded estimates of the neutrino that undergoes strong interactions:

Name Symbol Charge Rest mass


(electron charges) (MeV/c 2)

electron e 1 0.511
electron neutrino e 0 *
muon 1 105.7
muon neutrino 0 *
tauon 1 1784
tau neutrino 0 *

Table of leptons
283 elements of an orbit

baryons are composed of three quarks (u, quickly identied as the carrier of a fth
d, or s) or three antiquarks (, , or ); quark, bottom (b).
mesons are composed of (u, d, or s) Thus, by the end of 1977, ve avours
quarkantiquark pairs. of quark (u, d, s, c, b) were known to exist
No other combinations seemed to be together with six avours of lepton (e, ,
necessary to describe the full variation of , e, , ). Assuming that quarks and lep-
the observed hadrons. This scheme even tons are the fundamental constituents of
led to the prediction of other particles matter, many of the strong and weak in-
that were not known to exist in 1961. For teractions of hadrons and the weak inter-
example, in 1961 Gell-Mann not only pre- actions of leptons can be explained. e
dicted the (omega-minus) particle, but However, anticipating a symmetry in na-
more importantly told experimentalists tures building blocks, it was expected
exactly how to produce it. The particle that a sixth quark would eventually reveal
was nally discovered in 1964. itself. This quark, labelled top (t), would
Gell-Mann called his scheme the eight- be the 2/3 electronic charge partner to the
fold way, after the similarly named Bud- b quark (see table below).
dhist principle. The scheme requires that
quarks have properties not previously al- Quark symbol Name Charge
lowed for fundamental particles. For ex-
ample, quarks have fractional electric u up 2/3
charges, i.e. charges of 1/3 and 2/3 of the d down 1/3
electron charge. Quarks also have a strong c charm 2/3
afnity for each other through a new kind s strange 1/3
of charge known as colour charge. Thus t top 2/3
colour charge is responsible for strong b bottom 1/3
interactions, and the force is known as
the colour force. This is a revision of Table of quarks (mass is not shown because quarks
Yukawas proposal in 1930. Yukawas are never observed alone)
strong force was mediated by -mesons.
The strong force is now thought to be me-
In 1998 the top quark was found at
diated by exchange of particles carrying
CERN in Geneva and the symmetry of six
colour charge called gluons. The theory
quarks with six leptons was nally ver-
governing these colour charge combina-
ied. In 1978 the standard model was pro-
tions is modelled on *quantum electro-
posed as the denitive theory of the
dynamics and is known as *quantum
fundamental constituents of matter. In
chromodynamics.
the current view, all matter consists of
In November 1974 the discovery of the
three kinds of particles: leptons, quarks,
(psi) particle initiated what later came
and mediators (see table overleaf). The
to be known as the November revolu-
mediators are the particles by which the
tion. At the time, any known hadron
four fundamental interactions are medi-
could be described as some combination
ated. In the standard model, each of these
of u, d, or s quarks. These hadrons were
interactions has a particle mediator. For
very short-lived with lifetimes of about
the electromagnetic reaction it is the
1023 s. The particle, however, had a life-
*photon.
time of 1020 s; i.e. a thousand times
For weak interactions the force is medi-
longer. This suggested a completely differ-
ated by three particles called W+, W, and
ent species of particle. It is now univer-
Z *bosons; for the strong force it is the
sally accepted that the represents a
gluon. Current theories of quantum grav-
meson-bound state of a new fourth quark,
ity propose the *graviton as the mediator
the charm (c) quark and its antiquark. In
for the gravitational interaction, but this
1977 the list of quarks once again in-
work is highly speculative and the gravi-
creased with the discovery of a new even
ton has never been detected.
heavier meson, called the Y (upsilon)
meson. This meson was found to have an elements of an orbit Six parameters
even longer lifetime than the , and was used to dene the path of a celestial body.
elevation of boiling point 284

Interaction Mediator Rest mass Charge


(exchange particle) (GeV/c2)

strong gluon 0 0
electromagnetic photon 0 0
weak W+, W, Z 81,81,93 +1,1,0
gravitational graviton 0 0

e Table of mediators

The shape of the orbit is dened by its ec- sample are bound by the antibody. A sec-
centricity (see conic) and semimajor axis. ond antibody specic for a second site on
The orientation of the orbit is specied by the test protein is added; this is conju-
the *inclination of the orbital plane to the gated with an enzyme, which catalyses a
reference plane (usually the *ecliptic) and colour change in the fourth reagent,
by the longitude of the ascending *node added nally to the sheet. The colour
(the angular distance from the vernal change can be measured photometrically
equinox to the ascending node). The posi- and compared against a standard curve to
tion of the body in its orbit is dened by give the concentration of protein in the
its eccentric *anomaly and the position as sample. ELISA is widely used for diagnos-
a function of time is calculated from the tic and other purposes.
periapsis passage (see apsides).
ellipse A *conic formed by the intersec-
elevation of boiling point An in- tion of a plane with a right circular cone,
crease in the boiling point of a liquid so that the plane is inclined to the axis of
when a solid is dissolved in it. The eleva- the cone at an angle in excess of half the
tion is proportional to the number of par- apex angle of the cone. The ellipse has
ticles dissolved (molecules or ions) and is two vertices, which are joined by a line
given by t = kBC, where C is the molal called the major axis. The centre of the el-
concentration of solute. The constant kB is lipse falls on this line, midway between
the ebullioscopic constant of the solvent the vertices. The minor axis is the line per-
and if this is known, the molecular pendicular to the major axis that passes
weight of the solute can be calculated through the centre and joins two points
from the measured value of t. The eleva- on the ellipse. The foci of an ellipse are
tion is measured by a Beckmann ther- two points on the major axis so placed
mometer. See also colligative properties. that for any point on the ellipse the sum
of the distances from that point to each
elimination reaction A reaction in
focus is constant. (See illustration.) The
which one molecule decomposes into
area of an ellipse is ab, where a and b are
two, one much smaller than the other.
half the major and minor axes, respec-
Elinvar Trade name for a nickel tively. For an ellipse centred at the origin,
chromium steel containing about 36% the equation in Cartesian coordinates is
nickel, 12% chromium, and smaller x2/a2 + y2/b2 = 1. The foci are at (ea, 0) and
proportions of tungsten and manganese. (ea, 0), where e is the eccentricity. Each
Its elasticity does not vary with tempera- of the two chords of the ellipse passing
ture.
ELISA (enzyme-linked immunosorbent latus minor axis
rectum
assay) A sensitive technique (see immuno-
assay) for accurately determining the major axis
amount of protein or other antigen in a
given sample by means of an enzyme- centre
catalysed colour change. Antibody specic vertex vertex
focus focus
to the test protein is adsorbed onto a solid
substrate, such as a PVC sheet, and a
measured amount of the sample is added;
all molecules of the test protein in the An ellipse
285 embryo sac

through a focus and parallel to the minor Rn220 (thoron emanation), and Rn219
axis is called a latus rectum and has a (actinium emanation).
length equal to 2b2/a.
emasculation The removal of the an-
ellipsoid A solid body formed when an thers of a ower in order to prevent self-
*ellipse is rotated about an axis. If it is ro- pollination or the undesirable pollination
tated about its major axis it is a prolate el- of neighbouring plants.
lipsoid; if it is rotated about its minor axis
EmbdenMeyerhof pathway See gly-
it is an oblate ellipsoid. For an ellipsoid
colysis.
centred at the origin the equation in
Cartesian coordinates is: embryo 1. An animal in the earliest e
2 2 2 2 2 2 stages of its development, from the time
x /a + y /b + z /c = 1.
when the fertilized ovum starts to divide
(see cleavage), while it is contained
elliptical galaxy See galaxy.
within the egg or reproductive organs of
elliptical polarization See polariza- the mother, until hatching or birth. A
tion of light. human embryo (see illustration p. 288) is
called a *fetus after the rst eight weeks
El Nio A surge of warm ocean water
of pregnancy. 2. The structure in plants
(the Peru current) that occurs every 5 to 8
that develops from the zygote prior to
years off the eastern coast of South Amer-
germination. In seed plants the zygote is
ica. See Feature.
situated in the *embryo sac of the ovule.
Elton, Charles Sutherland (190091) It divides by mitosis to form the embry-
British zoologist and ecologist, who onic cell and a structure called the sus-
founded the Bureau of Animal Population pensor, which embeds the embryo in the
at Oxford in 1932 and the same year be- surrounding nutritive tissue. The embry-
came editor of the new Journal of Animal onic cell divides continuously and eventu-
Ecology. The rst zoologist to study ani- ally gives rise to the *radicle (young root),
mals in relation to their environment, he *plumule (young shoot), and one or two
explored the nature of food chains and *cotyledons (seed leaves). Changes also
studied population uctuations. take place in the surrounding tissues of
the ovule, which becomes the *seed en-
eluate See chromatography; elution.
closing the embryo plant.
eluent See chromatography; elution.
embryology The study of the develop-
elution The process of removing an ad- ment of animals from the fertilized egg to
sorbed material (adsorbate) from an ad- the new adult organism. It is sometimes
sorbent by washing it in a liquid (eluent). limited to the period between fertilization
The solution consisting of the adsorbate of the egg and hatching or birth (see em-
dissolved in the eluent is the eluate. bryo).
Elution is the process used to wash com-
embryophyte A true plant, i.e. one that
ponents of a mixture through a *chroma-
develops from an embryo and therefore is
tography column.
necessarily multicellular. The term under-
elutriation The process of suspending lines the distinction between plants and
nely divided particles in an upward ow- algae, which lack embryos.
ing stream of air or water to wash and
embryo sac A large cell that develops
separate them into sized fractions.
in the *ovule of owering plants. It is
elytra The thickened horny forewings of equivalent to the female *gametophyte of
the *Coleoptera (beetles), which cover lower plants, although it is very much re-
and protect the membranous hindwings duced. Typically, it contains eight nuclei
when the insect is at rest. formed by division of the original female
gamete (see illustration). One forms the
e-mail See electronic mail.
*oosphere (egg cell), which is fertilized by
emanation The former name for the a male nucleus and becomes the *em-
gas radon, of which there are three iso- bryo. The two polar nuclei fuse with a sec-
topes: Rn222 (radium emanation), ond male nucleus to form a triploid
286

EL NI O

A phenomenon reoccurring every few years in the equatorial part of the


Pacific ocean, characterized by movement of a mass of warm water eastward
towards the west coast of South America. This change in ocean conditions has
long been recognized in Peru, where sailors noticed that an unusual counter-
current appeared in certain years around the area of the port of Paita. They
named this current El Nio the Christ Child because it usually appeared
e immediately after Christmas. It was also known that the appearance of this
current coincided with different weather conditions, particularly increased
rainfall and sometimes flooding.
The phenomenon has dramatic effects on the climate and ecology of this
part of South America. In particular, it interrupts the Humbolf current, which
is a cold ocean current carrying plankton from Arctic regions. El Nio occurs
every 47 years and the effects last for about 8 months. In the 1960s interest
developed in the phenomenon and it is now recognized that El Nio has
effects on climate much wider than those observed on the west coast of South
America. In extreme cases, as in 198687 and 199798, it can cause tropical
cyclones over the whole Pacific area, drought in southeastern Asia and
Australia, and increased rainfall and flooding in parts of North America.
The mechanism of El Nio
There is no definite agreement about what induces an El Nio event, but the
physical mechanism of how it occurs is fairly well understood. The large-scale
movement of water in the worlds oceans is influenced by, and in turn
influences, the prevailing wind patterns. In tropical regions there are
persistent trade winds flowing from east to west. In the Pacific these push
large amounts of water westward towards the coasts of Indonesia. This causes
a significant difference in sea level between opposite sides of the Pacific. For
instance, the sea level in the Philippines in the west is around 60 cm higher
than that on the coast of Panama in the eastern Pacific. The water in the west
is also much warmer. This mass of warm ocean in the western Pacific gives
Indonesia its high rainfall under normal climatic conditions.

UNITED
STATES
normal trade winds

warm
normal ocean currents
PERU
INDONESIA

waters

AUSTRALIA

In normal years the trade winds blow from east to west across the Pacific. This causes a
difference in sea level and a mass of warm water builds up in the western Pacific, creating a
warm area of ocean off the east coast of Indonesia. This area has some of the highest rainfall
in the world.
287

Every few years there is a significant change in the prevailing wind pattern,
involving a fall in the intensity of the trade winds. In extreme cases, a reversal
of direction of the winds may occur. As a result, the warm water that is piled
up in the west flows back eastwards across the Pacific towards the west coast
of South America. The event lasts until the normal conditions and wind
patterns are re-established.
The Southern Oscillation and La Nia
El Nio events are connected with another periodic phenomenon occurring e
not in the ocean but in the atmosphere. The meteorologist Sir Gilbert Walker
noted in 1923 that when pressure was high in the Pacific it tends to be low in
the Indian ocean, and vice versa. The extent of this is now measured by taking
the difference between the surface atmospheric pressure at Darwin in
Australia and at Tahiti in the south Pacific. A high pressure at one site is
usually accompanied by a low pressure at the other and every few years the
pattern reverses. There is a large mass of air slowly oscillating (with a period of
a few years) across tropical regions. Walker called this the Southern Oscillation.
The Southern Oscillation is part of a large general cycle of coupled air and
water flow known as the El NioSouthern Oscillation (ENSO). El Nio is the
warm phase of this cycle. In some years, as part of the ENSO cycle, a cold
region develops in the eastern tropical Pacific. This is known as La Nia (the
little girl).
The Southern oscillation is not the only atmospheric oscillation that occurs.
For example there is a North Atlantic Oscillation (NAO) measured by the
pressure difference between Iceland and the Azores, which is thought to have
a major influence on climatic conditions and on the ecosystems of this part of
the globe. There is however no Atlantic analogue of El Nio.
The causes of El Nio
Although the flows of air and water are understood, there is no concensus
about why an El Nio forms. Some workers have suggested that the frequency
and intensities of El Nio events may be increasing because of global warming.
However, there is no direct evidence for this .

El Nio current

weakened
trade winds increased
surface
temperatures

In certain years there is a reduction in the intensity of the trade winds across the Pacific. This
allows the mass of warm water in the west to flow across to the east, creating a warm area
of ocean off the west coast of South America. This brings rain to Peru but causes drought in
southeast Asia and Australia.
emerald 288

chorionic villi uterus wall

fallopian tube placenta

blood vessel
yolk sac

e umbilical cord

amnion

embryo

chorion

cervix amniotic fluid

vagina

A developing human embryo

nucleus that gives rise to the *endosperm. e.m.f. See electromotive force.
The three remaining nuclei form the an-
emission spectrum See spectrum.
tipodal cells.
emissivity Symbol . The ratio of the
antipodal cells power per unit area radiated by a surface
to that radiated by a *black body at the
same temperature. A black body therefore
embryo sac has an emissivity of 1 and a perfect reec-
tor has an emissivity of 0. The emissivity
polar nuclei of a surface is equal to its *absorptance.
integuments emittance See exitance.
oosphere egg emitter See transistor.
synergids apparatus
emitter-coupled logic (ECL) A set of
integrated *logic circuits. The input part
Embryo sac of an ECL consists of an emitter-coupled
*transistor pair which is a very good dif-
emerald The green gem variety of ferential amplier. The output is through
*beryl: one of the most highly prized an *emitter follower. ECL circuits are very
gemstones. The nest specimens occur in rapid logic circuits.
the Muzo mines, Colombia. Other occur- emitter follower An amplifying circuit
rences include the Ural Mountains, the using a bipolar junction *transistor with a
Transvaal in South Africa, and Kaligunan *common-collector connection. The out-
in India. Emeralds can also be successfully put is taken from the emitter.
synthesized.
empirical Denoting a result that is ob-
emery A rock composed of corundum tained by experiment or observation
(natural aluminium oxide, Al2O3) with rather than from theory.
magnetite, haematite, or spinel. It occurs
empirical formula See formula.
on the island of Naxos (Greece) and in
Turkey. Emery is used as an abrasive and emulsication (in digestion) The break-
polishing material and in the manufac- down of fat globules in the duodenum
ture of certain concrete oors. into tiny droplets, which provides a larger
289 endodermis

surface area on which the enzyme pancre- species that is restricted to one or a few
atic *lipase can act to digest the fats into localities in its distribution. Endemic
fatty acids and glycerol. Emulsication is species are usually conned to islands and
assisted by the action of the bile salts (see are vulnerable to extinction. 2. Describ-
bile). ing a disease or a pest that is always pres-
ent in an area. For example, malaria is
emulsion A *colloid in which small par-
endemic in parts of Africa.
ticles of one liquid are dispersed in an-
other liquid. Usually emulsions involve a endergonic reaction A chemical reac-
dispersion of water in an oil or a disper-
sion of oil in water, and are stabilized by
tion in which energy is absorbed. Compare
exergonic reaction.
e
an emulsier. Commonly emulsiers are
substances, such as *detergents, that have
endocarp See pericarp.
lyophobic and lyophilic parts in their mol- endocrine gland (ductless gland) Any
ecules. Dietary fats are reduced to an gland in an animal that manufactures
emulsion in the duodenum to facilitate *hormones and secretes them directly
their subsequent digestion (see emulsifi- into the bloodstream to act at distant sites
cation). in the body (known as target organs or
enamel The material that forms a cover- cells). Endocrine glands tend to control
ing over the crown of a *tooth (i.e. the slow long-term activities in the body, such
part that projects above the gum). Enamel as growth and sexual development. In
is smooth, white, and extremely hard, mammals they include the *pituitary,
being rich in minerals containing cal- *adrenal, *thyroid, and *parathyroid
cium, especially *apatite. It is produced by glands, the *ovary and *testis, the *pla-
certain cells (ameloblasts) of the oral epi- centa, and part of the pancreas (see islets
thelium and protects the underlying den- of langerhans). The activity of endocrine
tine of the tooth. Enamel may also be glands is controlled by negative feedback,
found in the placoid *scales of certain i.e. a rise in output of hormone inhibits a
sh, which demonstrates the common de- further increase in its production, either
velopmental origin of scales and teeth. directly or indirectly via the target organ
or cell. See also neuroendocrine system.
enantiomers See optical activity. Compare exocrine gland.
enantiomorphism See optical activ- endocrinology The study of the struc-
ity. ture and functions of the *endocrine
enantiotropy See allotropy. glands and of the *hormones they pro-
duce.
encephalin (enkephalin) Any of a class
of *endorphins consisting of ve amino endocytosis The process by which ma-
acids and found principally in the brain. terials enter a cell without passing
They bind to opiate receptors in the brain through the plasma membrane. The
and their release controls levels of pain membrane folds around material outside
and other sensations. the cell, resulting in the formation of a
saclike vesicle into which the material is
endangered species A plant or animal incorporated. This vesicle is then pinched
species dened by the IUCN (International
off from the cell surface so that it lies
Union for the Conservation of Nature and
within the cell. Both *phagocytosis and
Natural Resources) as being in immediate
*pinocytosis are forms of endocytosis.
danger of *extinction because its popula-
Compare exocytosis.
tion numbers have reached a critical level
or its habitats have been drastically re- endoderm (entoderm) The internal
duced. If these causal factors continue the layer of cells of the *gastrula, which will
species is unlikely to survive. A list of en- develop into the alimentary canal (gut)
dangered species is published by the and digestive glands of the adult. See also
IUCN, which also denes other categories germ layers.
of threatened species.
endodermis The innermost layer of the
endemic 1. Describing a plant or animal root *cortex of a plant, lying immediately
endoergic 290

outside the vascular tissue. Various tween the plasma and nuclear mem-
modications of the endodermal cell walls branes and is the site of protein synthesis.
enable them to regulate the passage of It is also concerned with the transport of
materials both into and out of the vascu- proteins and lipids within the cell. Rough
lar system. An endodermis may also be ER has *ribosomes attached to its surface;
seen in the stems of some plants. proteins synthesized on the ribosomes are
enclosed in vesicles and transported to
endoergic Denoting a nuclear process
the *Golgi apparatus. Smooth ER lacks
that absorbs energy. Compare exoergic.
ribosomes; it is the site of important
e endogamy The fusion of reproductive metabolic reactions, including phospho-
cells from closely related parents, i.e. *in- lipid and fatty-acid synthesis.
breeding. Compare exogamy.
end organ The structure at the end of a
endogenous Describing a substance, peripheral nerve. Examples of end organs
stimulus, organ, etc., that originates from are the muscle *end plate at the end of a
within an organism. For example, growth motor neuron and the *receptor at the
rhythms not directed by environmental end of a sensory neuron.
stimuli are termed endogenous rhythms.
Lateral roots, which always grow from in- endorphin Any of a group of substances
side the main root rather than from its in the brain and other tissues that have
surface, are said to arise endogenously. pain-relieving properties similar to those
Compare exogenous. of morphine. They are all peptides or
polypeptides and include the *encepha-
endolymph The uid that lls the lins. Several endorphins are associated
membranous labyrinth of the vertebrate with the pituitary, while others occur in
*inner ear. See cochlea; semicircular the placenta, adrenal gland, pancreas, and
canals. Compare perilymph. various other tissues.
endometrium The mucous membrane endoskeleton A supporting framework
that lines the *uterus of mammals. It is that lies entirely within the body of an an-
comprised of an upper mucus-secreting imal, such as the bony *skeleton of verte-
layer, which is shed during menstruation, brates or the spicules of a sponge. The
and a basal layer, which proliferates to function of an endoskeleton is to support
form the upper layer. See also menstrual the body and in vertebrates it also pro-
cycle. tects the organs and provides a system of
endonuclease An enzyme that cataly- levers on which the muscles can act to
ses the internal cleavage of nucleic acids. produce movement. Compare exoskel-
See also restriction enzyme. Compare ex- eton.
onuclease. endosperm A nutritive tissue, charac-
endoparasite A parasite that lives in- teristic of owering plants, that surrounds
side its hosts body. See parasitism. the developing embryo in a seed. It devel-
ops from nuclei in the *embryo sac and
endopeptidase A protein-digesting en- its cells are triploid. In endospermic seeds
zyme that cleaves a polypeptide chain at it remains and increases in size; in non-
specic sites between amino acids. For ex- endospermic seeds it disappears as the
ample, *chymotrypsin cleaves the chain food is absorbed by the embryo, particu-
next to aromatic amino acids, such as larly the *cotyledons. Many plants with
phenylalanine; *trypsin cleaves the chain endospermic seeds, such as cereals and oil
next to basic amino acids, such as lysine
crops, are cultivated for the rich food re-
or arginine; and *pepsin cleaves the chain
serves in the endosperm.
next to tyrosine and phenylalanine. Com-
pare exopeptidase. endospore The resting stage of certain
bacteria, formed in response to adverse
endoplasm See cytoplasm.
conditions. The bacterial cell becomes en-
endoplasmic reticulum (ER) A system closed in a tough resistant protein coat.
of membranes within the cytoplasm of On return to favourable conditions the
plant and animal *cells. It forms a link be- spore germinates and reverts to the nor-
291 energy band

mal vegetative form of the organism. tion that takes heat from its surround-
Endospores can remain viable for long pe- ings. Compare exothermic.
riods, perhaps several thousands of years.
endotoxin See toxin.
endosymbiont theory A theory, de-
end plate The area of the plasma mem-
vised principally by US biologist Lynn
brane of a muscle cell that lies immedi-
Margulis (1938 ), that eukaryotic organ-
ately beneath a motor nerve ending at a
isms evolved from symbiotic associations
*neuromuscular junction. Release of a
between prokaryotic ancestors. Free-living
*neurotransmitter at the end plate in-
aerobic bacteria and chloroxybacteria (see
cyanobacteria) became incorporated in- duces contraction of the muscle bre. e
side larger nucleated prokaryotic cells, end point The point in a titration at
where they acted as forerunners of the which reaction is complete as shown by
mitochondria and chloroplasts seen in the *indicator.
modern eukaryotes. Such events are held
to have occurred on several occasions, energy A measure of a systems ability
producing various lineages of both to do work. Like work itself, it is meas-
heterotrophic and phototrophic protoc- ured in joules. Energy is conveniently clas-
tists, from which evolved ancestors of ani- sied into two forms: potential energy is
mals, plants, and fungi. There is strong the energy stored in a body or system as a
evidence for the theory, particularly the consequence of its position, shape, or
nding that mitochondria and chloro- state (this includes gravitational energy,
plasts have DNA similar in form to that electrical energy, nuclear energy, and
of eubacteria, and that they contain chemical energy); kinetic energy is energy
prokaryotic-type ribosomes. of motion and is usually dened as the
work that will be done by the body pos-
endothelium A single layer of thin sessing the energy when it is brought to
platelike cells that line the inner surfaces rest. For a body of mass m having a speed
of blood and lymph vessels and the heart. v, the kinetic energy is mv2/2 (classical) or
Endothelium is derived from the *meso- (m m0)c2 (relativistic). The rotational ki-
derm. Compare epithelium; mesothelium. netic energy of a body having an angular
endotherm (homoiotherm) An animal velocity is I2/2, where I is its moment
that can generate and maintain heat of inertia.
within its body independently of the envi- The *internal energy of a body is the
ronmental temperature. Mammals and sum of the potential energy and the ki-
birds are endotherms; they are often de- netic energy of its component atoms and
scribed as being warm-blooded. See molecules.
homoiothermy. Compare ectotherm.
energy band A range of energies that
endothermic Denoting a chemical reac- electrons can have in a solid. In a single

E E E
conduction
band

forbidden band conduction conduction band


band
electron
distribution
valence band
valence valence
band band

insulator conductor semiconductor

Energy bands
energy flow 292

atom, electrons exist in discrete *energy proportion of energy is lost at each


levels. In a crystal, in which large num- trophic level, mostly in the form of heat
bers of atoms are held closely together in energy due to respiration, a food chain
a lattice, electrons are inuenced by a does not normally consist of more than
number of adjacent nuclei and the ve trophic levels: the fth trophic level
sharply dened levels of the atoms be- does not contain enough energy to sup-
come bands of allowed energy (see illus- port further levels. Energy is also lost
tration); this approach to energy levels in from the food chain in excretory products
solids is often known as the band theory. and the remains of dead organisms; this is
e Each band represents a large number of converted into heat energy by the action
allowed quantum states. Between the of *decomposers. See also productivity;
bands are forbidden bands. The outermost pyramid of energy.
electrons of the atoms (i.e. the ones re-
energy level A denite xed energy
sponsible for chemical bonding) form the
that a system described by *quantum me-
valence band of the solid. This is the
chanics, such as a molecule, atom, elec-
band, of those occupied, that has the
tron, or nucleus, can have. In an atom, for
highest energy.
example, the atom has a xed energy cor-
The band structure of solids accounts
responding to the *orbitals in which its
for their electrical properties. In order to
electrons move around the nucleus. The
move through the solid, the electrons
atom can accept a quantum of energy to
have to change from one quantum state
become an excited atom (see excitation)
to another. This can only occur if there
if that extra energy will raise an electron
are empty quantum states with the same
to a permitted orbital. Between the
energy. In general, if the valence band is
ground state, which is the lowest possible
full, electrons cannot change to new
energy level for a particular system, and
quantum states in the same band. For con-
the rst excited state there are no permis-
duction to occur, the electrons have to be
sible energy levels. According to the
in an unlled band the conduction band.
*quantum theory, only certain energy lev-
Metals are good conductors either because
els are possible. An atom passes from one
the valence band and the conduction
energy level to the next without passing
band are only half-lled or because the
through fractions of that energy transi-
conduction band overlaps with the va-
tion. These levels are usually described by
lence band; in either case vacant states
the energies associated with the individ-
are available. In insulators the conduction
ual electrons in the atoms, which are al-
band and valence band are separated by a
ways lower than an arbitrary level for a
wide forbidden band and electrons do not
free electron. The energy levels of mol-
have enough energy to jump from one
ecules also involve quantized vibrational
to the other.
and rotational motion.
In intrinsic *semiconductors the forbid-
den gap is narrow and, at normal temper- Engels salt See potassium carbonate.
atures, electrons at the top of the valence
engine Any device for converting some
band can move by thermal agitation into
forms of energy into mechanical work. See
the conduction band (at absolute zero, a
heat engine; carnot cycle; internal-
semiconductor would act as an insulator).
combustion engine; steam engine.
Doped semiconductors have extra bands
in the forbidden gap. enols Compounds containing the group
CH=C(OH) in their molecules. See also
energy ow (in ecology) The ow of en-
ketoenol tautomerism.
ergy that occurs along a *food chain. En-
ergy enters the food chain at the level of enrichment The process of increasing
the *producers (usually plants) in the the abundance of a specied isotope in a
form of solar energy. The plants convert mixture of isotopes. It is usually applied
solar energy into chemical energy in the to an increase in the proportion of U235,
process of *photosynthesis. Chemical en- or the addition of Pu239 to natural ura-
ergy is passed from one trophic level to nium for use in a nuclear reactor or
the next through feeding. Since a large weapon.
293 enzyme

ensemble A set of systems of particles ergy (Q ) transferred to the system by heat


used in *statistical mechanics to describe divided by the thermodynamic tempera-
a single system. The concept of an ensem- ture (T) at which this occurs, i.e. S =
ble was put forward by the US scientist Q /T. However, all real processes are to a
Josiah Willard Gibbs (18391903) in 1902 certain extent irreversible changes and in
as a way of calculating the time average of any closed system an irreversible change
the single system, by averaging over the is always accompanied by an increase in
systems in the ensemble at a xed time. entropy.
An ensemble of systems is constructed In a wider sense entropy can be inter-
from knowledge of the single system and preted as a measure of disorder; the e
can be represented as a set of points in higher the entropy the greater the disor-
*phase space with each system of the en- der (see boltzmann formula). As any real
semble represented by a point. Ensembles change to a closed system tends towards
can be constructed both for isolated sys- higher entropy, and therefore higher dis-
tems and for open systems. order, it follows that the entropy of the
enterokinase (enteropeptidase) An en- universe (if it can be considered a closed
zyme in the small intestine that activates system) is increasing and its available en-
trypsinogen to *trypsin. ergy is decreasing (see heat death of the
universe). This increase in the entropy of
enteron (coelenteron; gastrovascular the universe is one way of stating the sec-
cavity) The body cavity of the coelenter- ond law of *thermodynamics.
ates, which has one opening functioning
both as mouth and anus. See cnidaria. environment (in ecology) The physical,
chemical, and biological conditions of the
enthalpy Symbol H. A thermodynamic
region in which an organism lives. See also
property of a system dened by H = U +
ecology; ecosystem.
pV, where H is the enthalpy, U is the inter-
nal energy of the system, p its pressure, environmental resistance The sum
and V its volume. In a chemical reaction total of the factors that prevent popula-
carried out in the atmosphere the pres- tions from continually growing and there-
sure remains constant and the enthalpy of fore tend to keep populations at constant
reaction, H, is equal to U + pV. For an levels. These factors include predators,
exothermic reaction H is taken to be disease, and a shortage of any of the vari-
negative. ous requirements for survival, such as
entoderm See endoderm. food, water, shelter, and light (which is
particularly important for plants). See also
entomology The study of insects. population growth.
entomophily Pollination of a ower in enzyme A protein that acts as a *cata-
which the pollen is carried on an insect. lyst in biochemical reactions. Each en-
Entomophilous owers are usually zyme is specic to a particular reaction or
brightly coloured and scented and often group of similar reactions. Many require
secrete nectar. In some species (e.g. prim- the association of certain nonprotein *co-
ulas) there are structural differences factors in order to function. The molecule
between the owers to ensure that cross- undergoing reaction (the substrate) binds
pollination occurs. Other examples of to a specic *active site on the enzyme
entomophilous owers are orchids and molecule to form a short-lived intermedi-
antirrhinums. Compare anemophily; hy- ate (see enzymesubstrate complex): this
drophily. greatly increases (by a factor of up to 1020)
entropy Symbol S. A measure of the un- the rate at which the reaction proceeds to
availability of a systems energy to do form the product. Enzyme activity is in-
work; in a closed system an increase in uenced by substrate concentration and
entropy is accompanied by a decrease in by temperature and pH, which must lie
energy availability. When a system under- within a certain range. Other molecules
goes a reversible change the entropy (S) may compete for the active site, causing
changes by an amount equal to the en- *inhibition of the enzyme or even irre-
enzyme inhibition 294

versible destruction of its catalytic proper- suggested, including the *lock-and-key


ties. mechanism.
Enzyme production is governed by a
Eocene The second geological epoch of
cells genes. Enzyme activity is further
the *Tertiary period. It extended from the
controlled by pH changes, alterations in
end of the Palaeocene epoch, about 54
the concentrations of essential cofactors,
million years ago, to the beginning of the
feedback inhibition by the products of the
Oligocene epoch, about 38 million years
reaction, and activation by another en-
ago. The term was rst proposed by Sir
zyme, either from a less active form or an
e inactive precursor (*zymogen). Such
Charles Lyell in 1833. In some classica-
tions of geological time the *Palaeocene is
changes may themselves be under the
included as part of the Eocene. Mammals
control of hormones or the nervous sys-
were dominant in the Eocene: rodents, ar-
tem. See also enzyme kinetics.
tiodactyls, carnivores, perissodactyls (in-
Enzymes are classied into six major
cluding early horses), and whales were
groups, according to the type of reaction
among the groups to make their rst ap-
they catalyse: (1) *oxidoreductases; (2)
pearance.
*transferases; (3) *hydrolases; (4) *lyases;
(5) *isomerases; (6) *ligases. The names of eosin One of a series of acidic dyes, used
most individual enzymes also end in -ase, in optical microscopy, that colours cyto-
which is added to the names of the sub- plasm pink and cellulose red. It is fre-
strates on which they act. Thus *lactase is quently used as a counterstain with
the enzyme that acts to break down lac- *haematoxylin for colouring tissue
tose; it is classied as a hydrolase. smears and sections of animal tissue.
enzyme inhibition See inhibition. ephemeral 1. (in botany) An *annual
plant that completes its life cycle in con-
enzyme kinetics The study of the rates
siderably less than one growing season. A
of enzyme-catalysed reactions. Rates of re-
number of generations can therefore
action are usually measured by using the
occur in one year. Many troublesome
puried enzyme in vitro with the substrate
weeds, such as groundsel and willowherb,
and then observing the formation of the
are ephemerals. Certain desert plants are
product or disappearance of the substrate.
also ephemerals, completing their life
As the concentration of the substrate is in-
cycles in a short period following rain.
creased the rate of reaction increases pro-
2. (in zoology) A short-lived animal, such
portionally up to a certain point, after
as a mayy.
which any further increase in substrate
concentration no longer increases the re- ephemeris A tabulation showing the
action rate (see michaelismenten curve). calculated future positions of the sun,
At this point, all active sites of the en- moon, and planets, together with other
zyme are saturated with substrate; any useful information for astronomers and
further increase in the rate of reaction navigators. It is published at regular inter-
will occur only if more enzyme is added. vals.
Reaction rates are also affected by the
ephemeris time (ET) A time system
presence of inhibitors (see inhibition),
that has a constant uniform rate as op-
temperature, and pH (see enzyme).
posed to other systems that depend on
enzyme-linked immunosorbent the earths rate of rotation, which has in-
assay See elisa. herent irregularities. It is reckoned from
an instant in 1900 (Jan 0d 12h) when the
enzymesubstrate complex The
suns mean longitude was 279.696 677 8.
intermediate formed when a substrate
The unit by which ephemeris time is
molecule interacts with the *active site of
measured is the tropical year, which con-
an enzyme. Following the formation of an
tains 31 556 925.9747 ephemeris seconds.
enzymesubstrate complex, the substrate
This fundamental denition of the *sec-
molecule undergoes a chemical reaction
ond was replaced in 1964 by the caesium
and is converted into a new product. Vari-
second of atomic time.
ous mechanisms for the formation of en-
zymesubstrate complexes have been epicalyx A ring of bracts below a ower
295 epiphysis

that resembles a calyx. It is seen, for ex- principally to protect the plant from in-
ample, in the strawberry ower. jury and to reduce water loss. Some epi-
dermal cells are modied to form guard
epicarp See pericarp.
cells (see stoma) or hairs of various types
epicentre The point on the surface of (see piliferous layer). In woody plants the
the earth directly above the focus of an functions of the shoot epidermis are
earthquake or directly above or below a taken over by the periderm tissues (see
nuclear explosion. cork cambium) and in mature roots the
epicotyl The region of a seedling stem epidermis is sloughed off and replaced by
above the stalks of the seed leaves (*coty- the *hypodermis. e
ledons) of an embryo plant. It grows epidiascope An optical instrument
rapidly in seeds showing *hypogeal ger- used by lecturers, etc., for projecting an
mination and lifts the stem above the soil enlarged image of either a translucent ob-
surface. Compare hypocotyl. ject (such as a slide or transparency) or an
epicycle A small circle whose centre opaque object (such as a diagram or
rolls around the circumference of a larger printed page) onto a screen.
xed circle. The curve traced out by a epididymis A long coiled tube in which
point on the epicycle is called an epicy- spermatozoa are stored in vertebrates. In
cloid. reptiles, birds, and mammals it is at-
epidemiology The study of diseases tached at one end to the *testis and opens
that affect large numbers of people. Tradi- into the sperm duct (*vas deferens) at the
tionally, epidemiologists have been con- other.
cerned primarily with infectious diseases, epigamic Serving to attract a mate.
such as typhoid and inuenza, that arise Epigamic characters include the bright
and spread rapidly among the population plumage of some male birds.
as epidemics. However, today the disci-
pline also covers noninfectious disorders, epigeal Describing seed germination in
such as diabetes, heart disease, and back which the seed leaves (cotyledons) emerge
pain. Typically the distribution of a dis- from the ground and function as true
ease is charted in order to discover pat- leaves. Examples of epigeal germination
terns that might yield clues about its are seen in sycamore and sunower. Com-
mode of transmission or the susceptibility pare hypogeal.
of certain groups of people. This in turn epiglottis A exible ap of cartilage in
may reveal insights about the causes of mammals that is attached to the wall of
the disease and possible preventive meas- the pharynx near the base of the tongue.
ures. During swallowing (see deglutition) it
epidermis 1. (in zoology) The outermost covers the *glottis (the opening to the res-
layer of cells of the body of an animal. In piratory tract) and helps to prevent food
invertebrates the epidermis is normally from entering the trachea (windpipe), al-
only one cell thick and is covered by an though it is not essential for this purpose.
impermeable *cuticle. In vertebrates the
epilimnion The upper layer of water in
epidermis is the thinner of the two layers
a lake. Compare hypolimnion. See thermo-
of *skin (compare dermis). It consists of a
cline.
basal layer of actively dividing cells (see
malpighian layer), covered by layers of epimerism A type of optical isomerism
cells that become impregnated with ker- in which a molecule has two chiral cen-
atin (see keratinization). The outermost tres; two optical isomers (epimers) differ
layers of epidermal cells (the *stratum in the arrangement about one of these
corneum) form a water-resistant protec- centres. See also optical activity.
tive layer. The epidermis may bear a
epinephrine See adrenaline.
variety of specialized structures (e.g.
*feathers, *hairs). 2. (in botany) The out- epiphysis The terminal section of a
ermost layer of cells covering a plant. It is growing bone (especially a long limb
overlaid by a *cuticle and its functions are bone) in mammals. It is separated from
epiphyte 296

the bone shaft (diaphysis) by cartilage. *endoderm. Compare endothelium;


New bone is produced on the side of the mesothelium.
cartilage facing the diaphysis, while new
epithermal neutron A neutron with
cartilage is produced on the other side of
an energy in excess of that associated
the cartilage disc. When the bone reaches with a thermal neutron (see moderator)
adult length the epiphysis merges with but less than that of a *fast neutron, i.e. a
the diaphysis. neutron having an energy in the range 0.1
epiphyte A plant that grows upon an- to 100 eV.
e other plant but is neither parasitic on it
nor rooted in the ground. Epiphytes in-
EPM See electron probe microanalysis.
clude many mosses and lichens and some epoxides Compounds that contain oxy-
tropical orchids. gen atoms in their molecules as part of a
three-membered ring (see formula). Epox-
episome A genetic element that can ides are thus cyclic ethers.
exist and replicate either independently
of its host cells chromosomes or as an in- _ _ R3
_ C___C _ R
R1
tegrated part of the chromosomes. Exam- R2
O 4
ples include certain bacterial *plasmids.
epistasis A gene interaction in which The functional group in epoxides
one gene suppresses the effect of another
gene that is situated at a different *locus epoxyethane (ethylene oxide) A colour-
on the chromosome. For example, in less ammable gas, C2H4O; m.p. 111C;
guinea pigs the gene that controls the b.p. 13.5C. It is a cyclic ether (see epox-
production of melanin is epistatic to the ides) that is made by the catalytic oxi-
gene that regulates the deposition of dation of ethene. It can be hydrolysed to
melanin. A dominant allele (C) is responsi- ethane-1,2-diol and also polymerizes to:
ble for the production of melanin, while
OC2H4OC2H4, which is used for
the amount of melanin deposited is con-
lowering the viscosity of water (e.g. in re
trolled by a second gene, which deter-
ghting).
mines whether the coat colour is black or
brown. If an animal is homozygous reces- epoxy resins Synthetic resins produced
sive (cc) for melanin production, the coat by copolymerizing epoxide compounds
colour will be white regardless of the al- with phenols. They contain O linkages
leles that produce black or brown colora- and epoxide groups and are usually vis-
tion. cous liquids. They can be hardened by ad-
epitaxy (epitaxial growth) Growth of a dition of agents, such as polyamines, that
layer of one substance on a single crystal form cross-linkages. Alternatively, cata-
of another, such that the crystal structure lysts may be used to induce further poly-
in the layer is the same as that in the sub- merization of the resin. Epoxy resins are
strate. It is used in making semiconductor used in electrical equipment and in the
devices. chemical industry (because of resistance
epithelium A tissue in vertebrates con- to chemical attack). They are also used as
sisting of closely packed cells in a sheet adhesives.
with little intercellular material. It covers epsomite A mineral form of *magne-
the outer surfaces of the body and the sium sulphate heptahydrate, MgSO4.
walls of the internal cavities (coeloms). It 7H2O.
also forms glands and parts of sense or-
gans. Its functions are protective, absorp- Epsom salt See magnesium sulphate.
tive, secretory, and sensory. The types of EpsteinBarr virus See herpesvirus.
cell vary, giving rise to squamous,
cuboidal, columnar, and ciliated epithelia. equation of motion (kinematic equa-
Stratied epithelium (e.g. in the skin) is tion) Any of four equations that apply to
made up of several layers of cells. Epi- bodies moving linearly with uniform ac-
thelium is derived from *ectoderm and celeration (a). The equations, which relate
297 equilibrium constant

distance covered (s) to the time taken (t), the inclination of the ecliptic to the celes-
are: tial equator.
v = u + at equator 1. The great circle around the
earth that lies in a plane perpendicular to
s = (u + v)t/2
the earths axis. It is equidistant from the
s = ut + at2/2 two geographical poles. 2. The magnetic
equator is a line of zero magnetic dip (see
v2 = u2 + 2as,
geomagnetism) that is close to the geo-
where u is the initial velocity of the body
and v is its nal velocity.
graphical equator but lies north of it in
Africa and south of it in America. 3. The
e
celestial equator is the circle formed on
equation of state An equation that re-
the *celestial sphere by the extension of
lates the pressure p, volume V, and ther-
the earths equatorial plane. 4. (in cell bi-
modynamic temperature T of an amount
ology) See spindle.
of substance n. The simplest is the ideal
*gas law: equilibrium A state in which a system
has its energy distributed in the statisti-
pV = nRT,
cally most probable manner; a state of a
where R is the universal gas constant. Ap- system in which forces, inuences, reac-
plying only to ideal gases, this equation tions, etc., balance each other out so that
takes no account of the volume occupied there is no net change.
by the gas molecules (according to this A body is in static equilibrium if the re-
law if the pressure is innitely great the sultants of all forces and all couples acting
volume becomes zero), nor does it take on it are both zero; it may be at rest and
into account any forces between mol- will certainly not be accelerated. Such a
ecules. A more accurate equation of state body at rest is in stable equilibrium if after
would therefore be a slight displacement it returns to its orig-
inal position for a body whose weight is
(p + k)(V nb) = nRT,
the only downward force this will be the
where k is a factor that reects the de- case if the vertical line through its centre
creased pressure on the walls of the con- of gravity always passes through its base.
tainer as a result of the attractive forces If a slight displacement causes the body to
between particles, and nb is the volume move to a new position, then the body is
occupied by the particles themselves in unstable equilibrium.
when the pressure is innitely high. In A body is said to be in thermal equilib-
the van der Waals equation of state, pro- rium if no net heat exchange is taking
posed by the Dutch physicist J. D. van der place within it or between it and its sur-
Waals (18371923), roundings. A system is in *chemical equi-
librium when a reaction and its reverse
k = n2a/V2,
are proceeding at equal rates (see also equi-
where a is a constant. This equation more librium constant). These are examples
accurately reects the behaviour of real of dynamic equilibrium, in which activity
gases; several others have done better but in one sense or direction is in aggregate
are more complicated. balanced by comparable reverse activity.
equation of time The length of time equilibrium constant For a reversible
that must be added to the mean solar reaction of the type
time, as shown on a clock, to give the ap-
xA + yB zC + wD
parent solar time, as shown by a sundial.
The amount varies during the year, being chemical equilibrium occurs when the
a minimum of 14.2 minutes in February rate of the forward reaction equals the
and a maximum of +16.4 minutes in Octo- rate of the back reaction, so that the con-
ber. It is zero on four days (April 15/16, centrations of products and reactants
June 14/15, Sept. 1/2, Dec. 25/26). The dif- reach steady-state values. It can be shown
ference arises as a result of two factors: that at equilibrium the ratio of concentra-
the eccentricity of the earths orbit and tions
equilibrium law 298

[C]z[D]w/[A]x[B]y namics; it can be shown that the standard


equilibrium constant K is given by
is a constant for a given reaction and xed exp(G/RT), where G is the standard
temperature, called the equilibrium con- Gibbs free energy change for the com-
stant Kc (where the c indicates concentra- plete reaction. Strictly, the expressions
tions have been used). Note that, by above for equilibrium constants are true
convention, the products on the right- only for ideal gases (pressure) or innite
hand side of the reaction are used on the dilution (concentration). For accurate
top line of the expression for equilibrium work *activities are used.
e constant. This form of the equilibrium
constant was originally introduced in equilibrium law See equilibrium con-
1863 by C. M. Guldberg and P. Waage stant.
using the law of *mass action. They de- equinox 1. Either of the two points on
rived the expression by taking the rate of the *celestial sphere at which the *eclip-
the forward reaction tic intersects the celestial equator. The
kf [A]x[B]y sun appears to cross the celestial equator
from south to north at the vernal equinox
and that of the back reaction and from north to south at the autumnal
kb[C]z[D]w equinox. 2. Either of the two instants at
which the centre of the sun appears to
Since the two rates are equal at equilib- cross the celestial equator. In the north-
rium, the equilibrium constant Kc is the ern hemisphere the vernal equinox occurs
ratio of the rate constants kf /kb. The princi- on or about March 21 and the autumnal
ple that the expression is a constant is equinox on or about Sept. 23. In the
known as the equilibrium law or law of southern hemisphere the dates are re-
chemical equilibrium. versed. See precession of the equinoxes.
The equilibrium constant shows the
position of equilibrium. A low value of Kc equipartition of energy The theory,
indicates that [C] and [D] are small com- proposed by Ludwig Boltzmann and given
pared to [A] and [B]; i.e. that the back reac- some theoretical support by James Clerk
tion predominates. It also indicates how Maxwell, that the energy of gas molecules
the equilibrium shifts if concentration in a large sample under thermal *equilib-
changes. For example, if [A] is increased rium is equally divided among their avail-
(by adding A) the equilibrium shifts to- able *degrees of freedom, the average
wards the right so that [C] and [D] in- energy for each degree of freedom being
crease, and Kc remains constant. kT/2, where k is the *Boltzmann constant
For gas reactions, partial pressures are and T is the thermodynamic temperature.
used rather than concentrations. The sym- The proposition is not generally true if
bol Kp is then used. Thus, in the example *quantum considerations are important,
above but is frequently a good approximation.

Kp = pCzpDw/pAxpBy equivalence point The point in a titra-


tion at which reaction is complete. See in-
It can be shown that, for a given reaction dicator.
Kp = Kc(RT), where is the difference in
stoichiometric coefcients for the reac- equivalent proportions See chemical
tion (i.e. z + w x y). Note that the units combination.
of Kp and Kc depend on the numbers of equivalent weight The mass of an el-
molecules appearing in the stoichiometric ement or compound that could combine
equation. The value of the equilibrium with or displace one gram of hydrogen
constant depends on the temperature. If (or eight grams of oxygen or 35.5 grams
the forward reaction is exothermic, the of chlorine) in a chemical reaction. The
equilibrium constant decreases as the equivalent weight represents the combin-
temperature rises; if endothermic it in- ing power of the substance. For an el-
creases (see also vant hoffs isochore). ement it is the relative atomic mass
The expression for the equilibrium con- divided by the valency. For a compound it
stant can also be obtained by thermody- depends on the reaction considered.
299 escape velocity

erbium Symbol Er. A soft silvery metal- the surface or embedded in *boulder clay.
lic element belonging to the *lan- They provide geologists with information
thanoids; a.n. 68; r.a.m. 167.26; r.d. 9.006 about the movement of ice sheets.
(20C); m.p. 1529C; b.p. 2863C. It occurs
erythroblast Any of the cells in the
in apatite, gadolinite, and xenotine from
*myeloid tissue of red bone marrow that
certain sources. There are six natural iso-
develop into erythrocytes (red blood cells).
topes, which are stable, and twelve arti-
Erythroblasts have a nucleus and are at
cial isotopes are known. It has been used
rst colourless, but ll with *haemoglobin
in alloys for nuclear technology as it is a
neutron absorber; it is being investigated
as they develop. In mammals the nucleus
disappears. e
for other potential uses. It was discovered
by Carl Mosander (17971858) in 1843. erythrocyte (red blood cell) The most
numerous type of blood cell, which con-
erecting prism A glass prism used in
tains the red pigment *haemoglobin and
optical instruments to convert an inverted
is responsible for oxygen transport. Mam-
image into an erect image, as in prismatic
malian erythrocytes are disc-shaped and
binoculars.
lack a nucleus; those of other vertebrates
erg A unit of work or energy used in the are oval and nucleated. In man the num-
c.g.s. system and dened as the work done ber of erythrocytes in the blood varies be-
by a force of 1 dyne when it acts through tween 4.5 and 5.5 million per cubic
a distance of 1 centimetre. 1 erg = 107 millimetre. They survive for about four
joule. months and are then destroyed in the
spleen and liver. See also erythroblast.
ergocalciferol See vitamin d.
Compare leucocyte.
ergonomics The study of the engineer-
Esaki diode See tunnel diode.
ing aspects of the relationship between
workers and their working environment. ESCA See photoelectron spectroscopy.
ergosphere The region immediately escapement A device in a clock or
around a *black hole. The holes rotation watch that controls the transmission of
drags the spacetime continuum round power from the spring or falling weight
with it, so that frames of reference are to the hands. It is usually based on a bal-
not stationary with reference to the re- ance wheel or pendulum. It thus allows
mainder of the universe. The ergospheres energy to enter the mechanism in order
outer boundary is called the stationary to move the hands round the face, over-
limit. come friction in the gear trains, and main-
tain the balance wheel or pendulum in
ergosterol A *sterol occurring in fungi,
continuous motion.
bacteria, algae, and plants. It is converted
into vitamin D2 by the action of ultravio- escape velocity The minimum speed
let light. needed by a space vehicle, rocket, etc., to
escape from the gravitational eld of the
erosion The wearing away of the land
earth, moon, or other celestial body. The
surface by natural agents that involves the
gravitational force between a rocket of
transport of rock debris. These natural
mass m and a celestial body of mass M and
agents include moving waters (e.g. rivers,
radius r is MmG/r2 (see newtons law of
ocean waves), ice (e.g. glaciers), wind, or-
gravitation). Therefore the gravitational
ganisms, and gravity. See also soil
potential energy of the rocket with re-
erosion.
spect to its possible position very far from
erratic A fragment of rock, often unlike the celestial body on which it is resting
the rocks around it, that has been dis- can be shown to be GmM/r, assuming (by
placed from its original location by the convention) that the potential energy is
action of a glacier or, more rarely, an ice- zero at an innite distance from the celes-
berg. Erratics may have been moved as lit- tial body. If the rocket is to escape from
tle as several metres to more than 800 the gravitational eld it must have a ki-
km. They vary in size from small pebbles netic energy that exceeds this potential
to massive boulders, and may be found on energy, i.e. the kinetic energy mv2/2 must
Escherichia coli 300

be greater than MmG/r, or v > (2MG/r). range of elements in inorganic forms in


This is the value of the escape velocity. In- varying amounts, depending on the type
serting numerical values for the earth and of organism. The major elements, present
moon into this relationship gives an es- in tissues in relatively large amounts
cape velocity from the earth of 11 200 (greater than 0.005%), are calcium, phos-
m s1 and from the moon of 2370 m s1. phorus, potassium, sodium, chlorine,
sulphur, and magnesium (see also macro-
Escherichia coli (E. coli ) A species of
nutrient). The trace elements occur
Gram-negative aerobic bacteria that is
at much lower concentrations and thus
e found in the intestine and is also widely
used in microbiological and genetics re-
requirements are much less. The most
important are iron, manganese, zinc, cop-
search. The motile rod-shaped cells fer-
per, iodine, cobalt, selenium, molybde-
ment lactose and are usually harmless
num, chromium, and silicon (see also
commensals, although certain strains are
micronutrient). Each element may full
pathogenic and can cause a severe form of
one or more of a variety of metabolic
food poisoning. Studies of E. coli laboratory
roles. Sodium, potassium, and chloride
cultures have revealed much about the ge-
ions are the chief electrolytic components
netics of prokaryotes; the species is also
of cells and body uids and thus deter-
frequently used in genetic engineering,
mine their electrical and osmotic status.
particularly as a host for *gene cloning
Calcium, phosphorus, and magnesium are
and the expression of recombinant for-
all present in bone. Calcium is also essen-
eign genes in culture.
tial to nerve and muscle activity, while
esker An elongated, steep-sided ridge of phosphorus is a key constituent of the
debris left behind by meltwater streams chemical energy carriers (e.g. *ATP) and
owing in or under a slow-moving glacier. the nucleic acids. Sulphur is needed pri-
The ridge may be straight or, more often, marily for amino acid synthesis (in plants
sinuous in shape. It may be up to 50 m and microorganisms). The trace elements
high and 700 m wide. In very cold re- may serve as *cofactors or as constituents
gions, some eskers have ice cores. The de- of complex molecules, e.g. iron in haem
bris is stratied in layers and generally and cobalt in vitamin B12. See also mineral
consists of rounded particles of gravel and deficiency.
sand, with some ne-grain deposits. See
essential fatty acids *Fatty acids that
also kame.
must normally be present in the diet of
ESR See electron-spin resonance. certain animals, including man. Essential
fatty acids, which include *linoleic and
essential amino acid An *amino acid
*linolenic acids, all possess double bonds
that an organism is unable to synthesize
at the same two positions along their
in sufcient quantities. It must therefore
hydrocarbon chain and so can act as pre-
be present in the diet. In man the essen-
cursors of *prostaglandins. Deciency can
tial amino acids are arginine, histidine, ly-
cause dermatosis, weight loss, irregular
sine, threonine, methionine, isoleucine,
oestrus, etc. An adult human requires
leucine, valine, phenylalanine, and trypto-
210 g linoleic acid or its equivalent per
phan. These are required for protein syn-
day.
thesis and deciency leads to retarded
growth and other symptoms. Most of the essential oil A natural oil with a dis-
amino acids required by man are also es- tinctive scent secreted by the glands of
sential for all other multicellular animals certain aromatic plants. *Terpenes are the
and for most protozoans. main constituents. Essential oils are ex-
tracted from plants by steam distillation,
essential element Any of a number of
extraction with cold neutral fats or sol-
elements required by living organisms to
vents (e.g. alcohol), or pressing and used
ensure normal growth, development, and
in perfumes, avourings, and medicine.
maintenance. Apart from the elements
Examples are citrus oils, ower oils (e.g.
found in organic compounds (i.e. carbon,
rose, jasmine), and oil of cloves.
hydrogen, oxygen, and nitrogen), plants,
animals, and microorganisms all require a esterication A reaction of an alcohol
301 ethanoic acid

with an acid to produce an ester and It is used as a solid fuel in portable stoves
water; e.g. and in slug pellets.
CH3OH + C6H5COOH CH3OOCC6H5 + ethanamide (acetamide) A colourless
H2 O solid crystallizing in the form of long
white crystals with a characteristic smell
The reaction is an equilibrium and is nor-
of mice, CH3CONH2; r.d. 1.159; m.p.
mally slow, but can be speeded up by ad-
82.3C; b.p. 221.25C. It is made by the de-
dition of a strong acid catalyst. The ester
hydration of ammonium ethanoate or by
can often be distilled off so that the reac-
tion can proceed to completion. The re-
the action of ammonia on ethanoyl chlo-
ride, ethanoic anhydride, or ethyl
e
verse reaction is ester hydrolysis or
ethanoate.
*saponication. See also labelling.
ethane A colourless ammable gaseous
esters Organic compounds formed by hydrocarbon, C2H6; m.p. 183C; b.p.
reaction between alcohols and acids (see 89C. It is the second member of the
illustration). Esters formed from car- *alkane series of hydrocarbons and occurs
boxylic acids have the general formula in natural gas.
RCOOR. Examples are ethyl ethanoate,
CH3COOC2H5, and methyl propanoate, ethanedioic acid See oxalic acid.
C2H5COOCH3. Esters containing simple ethane-1,2-diol (ethylene glycol; glycol)
hydrocarbon groups are volatile fragrant A colourless viscous hygroscopic liquid,
substances used as avourings in the food CH2OHCH2OH; m.p. 11.5C; b.p. 198C. It
industry. Triesters, molecules containing is made by hydrolysis of epoxyethane
three ester groups, occur in nature as oils (from ethene) and used as an antifreeze
and fats. See also glyceride. and a raw material for making *polyesters
etaerio A cluster of fruits formed from (e.g. Terylene).
the unfused carpels of a single ower. For ethanoate (acetate) A salt or ester of
example, the anemone has an etaerio of ethanoic acid (acetic acid).
*achenes, larkspur an etaerio of *follicles,
and blackberry an etaerio of *drupes. ethanoic acid (acetic acid) A clear vis-
cous liquid or glassy solid *carboxylic
ethanal (acetaldehyde) A colourless acid, CH3COOH, with a characteristically
highly ammable liquid aldehyde, sharp odour of vinegar; r.d. 1.049; m.p.
CH3CHO; r.d. 0.78; m.p. 121C; b.p. 16.6C; b.p. 117.9C. The pure compound
20.8C. It is made from ethene by the is called glacial ethanoic acid. It is manu-
*Wacker process and used as a starting factured by the oxidation of ethanol or by
material for making many organic com- the oxidation of butane in the presence of
pounds. The compound polymerizes if di- dissolved manganese(II) or cobalt(II)
lute acid is added to give ethanal trimer ethanoates at 200C, and is used in mak-
(or paraldehyde), which contains a six- ing ethanoic anhydride for producing cel-
membered ring of alternating carbon and lulose ethanoates. It is also used in
oxygen atoms with a hydrogen atom and making ethenyl ethanoate (for polyvinyl-
a methyl group attached to each carbon acetate). The compound is formed by the
atom. It is used as a drug for inducing fermentation of alcohol and is present in
sleep. Addition of dilute acid below 0C vinegar, which is made by fermenting
gives ethanal tetramer (or metaldehyde), beer or wine. Vinegar made from
which has a similar structure to the ethanoic acid with added colouring mat-
trimer but with an eight-membered ring. ter is called nonbrewed condiment. In

O
C C2H5 O
CH3 O H H O C C2H5 + H2O
CH3 O
methanol propanoic acid
methyl propanoate water
Ester formation
ethanoic anhydride 302

living organisms it combines with *coen- ethanoyl group (acetyl group) The or-
zyme A to form acetyl coenzyme A, which ganic group CH3CO.
plays a crucial role in energy metabolism.
ethene (ethylene) A colourless am-
ethanoic anhydride (acetic anhydride) mable gaseous hydrocarbon, C2H4; m.p.
A pungent-smelling colourless liquid, 169C; b.p. 103.7C. It is the rst mem-
(CH3CO)2O, b.p. 139.5C. It is used in or- ber of the *alkene series of hydrocarbons.
ganic synthesis as an *ethanoylating It is made by cracking hydrocarbons from
agent (attacking an OH or NH group) petroleum and is now a major raw ma-
e and in the manufacture of aspirin and cel-
lulose plastics. It hydrolyses in water to
terial for making other organic chemicals
(e.g. ethanal, ethanol, ethane-1,2-diol). It
give ethanoic acid. can be polymerized to *polyethene. It oc-
curs naturally in plants, in which it acts as
ethanol (ethyl alcohol) A colourless
a *growth substance: its best-known ef-
water-soluble *alcohol, C2H5OH; r.d. 0.789
fect is the stimulation of fruit ripening.
(0C); m.p. 117.3C; b.p. 78.3C. It is the
active principle in intoxicating drinks, in ethenone See ketene.
which it is produced by fermentation of
ethenyl ethanoate (vinyl acetate)
sugar using yeast
An unsaturated organic ester,
C6H12O6 2C2H5OH + 2CO2 CH2:CHOOCCH3; r.d. 0.9; m.p. 100C;
b.p. 73C. It is made by catalytic reaction
The ethanol produced kills the yeast and
of ethanoic acid and ethene and used to
fermentation alone cannot produce
make polyvinylacetate.
ethanol solutions containing more than
15% ethanol by volume. Distillation can ether 1. (aether) A hypothetical
produce a constant-boiling mixture con- medium once believed to be necessary to
taining 95.6% ethanol and 4.4% water. support the propagation of electromag-
Pure ethanol (absolute alcohol) is made by netic radiation. It is now regarded as
removing this water by means of drying unnecessary and in modern theory elec-
agents. tromagnetic radiation can be propagated
The main industrial use of ethanol is as through empty space. The existence of
a solvent although at one time it was a the ether was rst called into question as
major starting point for making other a result of the *MichelsonMorley experi-
chemicals. For this it was produced by fer- ment. 2. See ethoxyethane; ethers.
mentation of molasses. Now ethene has
ethers Organic compounds containing
replaced ethanol as a raw material and in-
the group O in their molecules. Exam-
dustrial ethanol is made by hydrolysis of
ples are dimethyl ether, CH3OCH3, and
ethene.
diethyl ether, C2H5OC2H5 (see ethoxy-
ethanoylating agent (acetylating ethane). They are volatile highly am-
agent) A chemical reagent used to intro- mable compounds made by dehydrating
duce an ethanoyl group (COCH3) instead alcohols using sulphuric acid.
of hydrogen in an organic compound. Ex-
ethology The study of the biology of
amples include *ethanoic anhydride and
*animal behaviour. Central to the etholo-
ethanoyl chloride (acetyl chloride,
gists approach is the principle that
CH3COCl).
animal behaviour (like physical char-
ethanoyl chloride (acetyl chloride) A acteristics) is subject to evolution through
colourless liquid acyl chloride (see acyl natural selection. Ethologists therefore
halides), CH3COCl, with a pungent smell; seek to explain how the behaviour of an
r.d. 1.105; m.p. 112.15C; b.p. 50.9C. It is animal in its natural environment may
made by reacting ethanoic acid with a contribute to the survival of the maxi-
halogenating agent such as phos- mum number of its relatives and off-
phorus(III) chloride, phosphorus(V) chlo- spring. This involves recognizing the
ride, or sulphur dichloride oxide and is stimuli that are important in nature (see
used to introduce ethanoyl groups into or- sign stimulus) and how innate predispo-
ganic compounds containing OH, NH2, sitions interact with *learning in the de-
and SH groups. See acylation. velopment of behaviour (see instinct).
303 eudiometer

Studies of this sort were pioneered by gas, C2H2, with a characteristic sweet
Konrad Lorenz and Niko Tinbergen. odour; r.d. 0.618; m.p. 80.8C; b.p.
84.0C. It is the simplest member of the
ethoxyethane (diethyl ether; ether) A
*alkyne series of unsaturated hydrocar-
colourless ammable volatile *ether,
bons, and is prepared by the action of
C2H5OC2H5; r.d. 0.71; m.p. 116C; b.p.
water on calcium dicarbide or by adding
34.5C. It can be made by *Williamsons
alcoholic potassium hydroxide to 1,2-di-
synthesis. It is an anaesthetic and useful
bromoethane. It can be manufactured by
organic solvent.
heating methane to 1500C in the pres-
ethyl 3-oxobutanoate (ethyl aceto- ence of a catalyst. It is used in oxyacety- e
acetonate) A colourless liquid ester with lene welding and in the manufacture of
a pleasant odour, CH3COCH2COOC2H5; ethanal and ethanoic acid. Ethyne can be
r.d. 1.03; m.p. <80C; b.p. 180.4C. It can polymerized easily at high temperatures
be prepared by reacting ethyl ethanoate to give a range of products. The inorganic
(CH3COOC2H5) with sodium or sodium saltlike dicarbides contain the ion C22, al-
ethoxide. The compound shows ketoenol though ethyne itself is a neutral com-
*tautomerism and contains about 7% of pound (i.e. not a protonic acid).
the enol form, CH3C(OH):CHCOOC2H5, etiolation The abnormal form of
under normal conditions. Sometimes growth observed when plants grow in
known as acetoacetic ester, it is used in darkness or severely reduced light. Such
organic synthesis. plants characteristically have blanched
ethyl acetate See ethyl ethanoate. leaves and shoots, excessively long shoots,
and reduced leaves and root systems.
ethyl acetoacetonate See ethyl
3-oxobutanoate. Eubacteria See bacteria.

ethyl alcohol See ethanol. eucaryote See eukaryote.

ethylamine A colourless ammable euchromatin See chromatin.


volatile liquid, C2H5NH2; r.d. 0.69; m.p. Euclid (c. 330c. 260 bc) Greek mathe-
81C; b.p. 16.6C. It is a primary amine matician, who worked at the academy in
made by reacting chloroethane with am- Alexandria. He is best known for his 13-
monia and used in making dyes. volume work the Elements, which deals
ethylbenzene A colourless ammable mainly with geometry and established the
liquid, C6H5C2H5; r.d. 0.867; m.p. 95C; axiomatic method of reasoning.
b.p. 136C. It is made from ethene and Euclidean geometry The form of ge-
ethybenzene by a *FriedelCrafts reaction ometry set forth in the 13 volumes, called
and is used in making phenylethene (for the Elements, by Euclid. It is based on a se-
polystyrene). ries of axioms and applies to plane
gures. Compare non-euclidean geom-
ethyl bromide See bromoethane.
etry.
ethylene See ethene.
eudicot Any dicotyledonous owering
ethylene glycol See ethane-1,2-diol. plant whose pollen has three apertures
(i.e. triaperturate pollen), through one of
ethylene oxide See epoxyethane.
which the pollen tube emerges during
ethyl ethanoate (ethyl acetate) A pollination. Eudicots contrast with the so-
colourless ammable liquid ester, called primitive dicots, such as the mag-
C2H5OOCCH3; r.d. 0.9; m.p. 83.6C; b.p. nolia family, which have uniaperturate
77.06C. It is used as a solvent and in pollen (i.e. with a single aperture). Recent
avourings and perfumery. studies in molecular systematics have sug-
gested that the primitive dicots are more
ethyl group The organic group
closely related to the monocots than to
CH3CH2.
eudicots.
ethyl iodide See iodoethane.
eudiometer An apparatus for measur-
ethyne (acetylene) A colourless unstable ing changes in volume of gases during
eugenics 304

chemical reactions. A simple example is a 1 m to about 30 m in lakes and coastal


graduated glass tube sealed at one end waters, and rarely reaches depths of more
and inverted in mercury. Wires passing than 200 m in the open ocean. Compare
into the tube allow the gas mixture to be aphotic zone.
sparked to initiate the reaction between
europium Symbol Eu. A soft silvery
gases in the tube.
metallic element belonging to the *lan-
eugenics The study of methods of im- thanoids; a.n. 63; r.a.m. 151.96; r.d. 5.245
proving the quality of human populations (20C); m.p. 822C; b.p. 1597C. It occurs
e by the application of genetic principles.
Positive eugenics would seek to do this by
in small quantities in bastanite and mona-
zite. Two stable isotopes occur naturally:
selective breeding programmes. Negative europium151 and europium153, both of
eugenics aims to eliminate harmful genes which are neutron absorbers. Experimen-
(e.g. those causing haemophilia and col- tal europium alloys have been tried for
our blindness) by counselling any prospec- nuclear-reactor parts but until recently
tive parents who are likely to be *carriers. the metal has not been available in suf-
Euglenida A class of mostly unicellular cient quantities. It is widely used in the
protoctists (including Euglena) that move form of the oxide in phosphors for televi-
by means of undulipodia (agella). Most sion screens. It was discovered by Sir
euglenids are photosynthetic and inhabit William Crookes in 1889.
fresh water, and in some classication sys- Eustachian tube The tube that con-
tems are regarded as green algae (phylum nects the *middle ear to the back of the
Chlorophyta). However, they lack a cell throat (pharynx) in vertebrates. It is nor-
wall, being covered with a proteinaceous mally closed, but during swallowing it
*pellicle, and some forms are colourless opens to allow air into the middle ear,
and thus ingest food, since they cannot which equalizes the pressure on each side
photosynthesize. Euglenids are sometimes of the *tympanum (eardrum). It was
classied as a phylum, Euglenophyta, but named after the Italian anatomist Bar-
they are now more usually included in a tolomeo Eustachio (?152074).
larger phylum, Discomitochondria, to-
gether with three other groups of or- eustasy Fluctuations in sea level on a
ganisms, on the basis of a common global scale. One cause is the growth and
mitochondrial structure (characterized by decay of ice masses, such as glaciers and
disc-shaped cristae) and the absence of ice sheets (glacio-eustasy); since the last
sexual reproduction. ice age sea levels have gradually risen.
Tectonic movements can also change the
Eukarya See domain; eukaryote. volume of ocean basins (e.g. through sea-
eukaryote (eucaryote) An organism oor spreading). Global average sea level
consisting of cells in which the genetic rose between 0.1 and 0.2 m during the
material is contained within a distinct nu- 20th century.
cleus. All organisms except bacteria are eutectic mixture A solid solution con-
eukaryotes. Some authorities classify all sisting of two or more substances and
eukaryotes in the Eukarya, one of the having the lowest freezing point of any
three *domains of life. See cell. Compare possible mixture of these components.
prokaryote. The minimum freezing point for a set of
Eumetazoa See metazoa. components is called the eutectic point.
Low-melting-point alloys are usually eu-
euphotic zone (epipelagic zone; photic
tectic mixtures.
zone) The topmost layer of a lake or sea in
which there is sufcient light for net pri- euthanasia The act of ending the life of
mary production, i.e. where the energy a person or animal in order to prevent
xed by photosynthesis exceeds that lost further suffering, e.g. from an incurable
by respiration. The depth varies, depend- and painful disease. This can be achieved
ing on such factors as turbidity, supply of by administering a lethal drug or by with-
nutrients in the water, tidal turbulence, holding vital treatment. In human medi-
and temperature. It typically ranges from cine euthanasia is fraught with ethical
305 exa-

and legal problems, and is illegal in most containing an even number of protons
countries. Where it is practised, strict and an odd number of neutrons.
safeguards are enforced to ensure that the
event horizon See black hole.
patients wishes are determined and ad-
hered to. Euthanasia is widely performed evergreen (Describing) a plant that
in veterinary medicine. bears leaves throughout the year, each
leaf being shed independently of the oth-
Eutheria (Placentalia) An infraclass of
mammals in which the embryos are re- ers after two or three years. The leaves of
evergreens are often reduced or adapted
tained in a uterus in the mothers body
and nourished by a *placenta. The young in some way to prevent excessive water e
are thus fully protected during their em- loss; examples are the needles of conifers
bryonic development and kept at a con- and the leathery waxy leaves of holly.
stant temperature. Placental mammals Compare deciduous.
evolved during the Cretaceous period evocation The ability of experimental
(about 100 million years ago). Modern pla- stimuli (e.g. chemicals or tissue implants)
centals are a highly diverse group that oc- to cause unspecialized embryonic tissue
cupy all types of habitat in all parts of the to develop into specialized tissue.
world. They include the orders *Artio-
dactyla, *Carnivora, *Cetacea, *Chi- evolute The locus of the centres of cur-
roptera, *Insectivora, *Perissodactyla, vature of all the points on a given curve
*Primates, *Proboscidea, and *Rodentia. (called the involute).
Compare metatheria; prototheria. evolution The gradual process by which
eutrophic Describing a body of water the present diversity of plant and animal
(e.g. a lake) with an abundant supply of life arose from the earliest and most prim-
nutrients and a high rate of formation of itive organisms, which is believed to have
organic matter by photosynthesis. Pollu- been continuing for at least the past 3000
tion of a lake by *sewage or *fertilizers million years. Until the middle of the 18th
renders it eutrophic (a process called eu- century it was generally believed that
trophication). This stimulates excessive each species was divinely created and
growth of algae (see algal bloom); the xed in its form throughout its existence
death and subsequent decomposition of (see special creation). Lamarck was the
these increases the *biochemical oxygen rst biologist to publish a theory to ex-
demand and thus depletes the oxygen plain how one species could have evolved
content of the lake, resulting in the death into another (see lamarckism), but it was
of the lakes sh and other animals. Com- not until the publication of Darwins On
pare dystrophic; oligotrophic. the Origin of Species in 1859 that special cre-
evaporation The change of state of a ation was seriously challenged. Unlike
liquid into a vapour at a temperature Lamarck, Darwin proposed a feasible
below the boiling point of the liquid. mechanism for evolution and backed it up
Evaporation occurs at the surface of a liq- with evidence from the fossil record and
uid, some of those molecules with the studies of comparative anatomy and em-
highest kinetic energies escaping into the bryology (see darwinism; natural selec-
gas phase. The result is a fall in the aver- tion). The modern version of Darwinism,
age kinetic energy of the molecules of the which incorporates discoveries in genetics
liquid and consequently a fall in its tem- made since Darwins time, remains the
perature. most acceptable theory of species evolu-
tion (see also punctuated equilibrium).
evaporative cooling Cooling of a sub- More controversial, however, and still to
stance as a result of evaporation. See also be rmly claried, are the relationships
laser cooling. and evolution of groups above the species
eveneven nucleus An atomic nucleus level.
containing an even number of protons
exa- Symbol E. A prex used in the met-
and an even number of neutrons.
ric system to denote 1018 times. For exam-
evenodd nucleus An atomic nucleus ple, 1018 metres = 1 exametre (Em).
exbi- 306

exbi- See binary prefixes. *neurotransmitter (such as acetylcholine),


released from the presynaptic membrane,
excess electron An electron in a *semi-
binds to the postsynaptic membrane.
conductor that is not required in the
This will induce an *action potential in
bonding system of the crystal lattice and
the receiving neuron if the EPSP is large
has been donated by an impurity atom. It
enough. Compare inhibitory postsynap-
is available for conduction (excess conduc-
tic potential.
tion).
exciton An electronhole pair in a crys-
exchange force 1. A force resulting
e from the continued interchange of parti-
tal that is bound in a manner analogous
to the electron and proton of a hydrogen
cles in a manner that bonds their hosts to-
atom. It behaves like an atomic excitation
gether. Examples are the covalent bond
that passes from one atom to another and
involving electrons, and the strong inter-
may be long-lived. Exciton behaviour in
action (see fundamental interactions) in
*semiconductors is important.
which mesons are exchanged between nu-
cleons or gluons are exchanged between exclusion principle See pauli exclu-
quarks (see elementary particles). 2. See sion principle.
magnetism.
excretion The elimination by an organ-
excimer See exciplex. ism of the waste products that arise as a
result of metabolic activity. These prod-
exciplex A combination of two different
ucts include water, carbon dioxide, and
atoms that exists only in an excited state.
nitrogenous compounds. Excretion plays
When an exciplex emits a photon of elec-
an important role in maintaining the con-
tromagnetic radiation, it immediately dis-
stancy of an organisms *internal environ-
sociates into the atoms, rather than
ment (see homeostasis). In plants and
reverting to the ground state. A similar
simple animals waste products are ex-
transient excited association of two atoms
creted by simple diffusion from the body,
of the same kind is an excimer. An exam-
but higher animals have specialized or-
ple of an exciplex is the species XeCl* (the
gans and organ systems devoted to this
asterisk indicates an excited state), which
function. Examples of excretory organs in
can be formed by an electric discharge in
vertebrates are the lungs (for carbon diox-
xenon and chlorine. This is used in the ex-
ide and water), and the *kidneys (for ni-
ciplex laser, in which a population inver-
trogenous compounds (urea) and water).
sion is produced by an electrical
In addition, mammals excrete small
discharge.
amounts of urea, salts, and water from
excitation 1. A process in which a nu- the skin in sweat.
cleus, electron, atom, ion, or molecule ac-
exercise Increased muscular activity,
quires energy that raises it to a quantum
which results in an increase in metabolic
state (excited state) higher than that of its
rate, heart rate, and oxygen uptake. Exer-
*ground state. The difference between the
cise also causes an increase in *anaerobic
energy in the ground state and that in the
respiration in order to compensate for the
excited state is called the excitation en-
*oxygen debt, which results in a build-up
ergy. See collective excitation; energy
of lactic acid in the tissues.
level; quasiparticle. 2. The process of
applying current to the winding of an exergonic reaction A chemical reac-
electromagnet, as in an electric motor. tion in which energy is released (compare
3. The process of applying a signal to the endergonic reaction). An *exothermic
base of a transistor or the control elec- reaction is an exergonic reaction in which
trode of a thermionic valve. energy is released in the form of heat.
excitatory postsynaptic potential exfoliation A type of rock erosion that
(EPSP) The electric potential that is gener- results from weathering. Layers or shells
ated in a postsynaptic neuron during the of rock are gradually removed from a
transmission of a nerve impulse (see massive outcrop, causing so-called onion-
synapse). It is caused by *depolarization skin weathering. Commonly affecting
of the postsynaptic membrane when a basalt and granite, it is thought to be
307 expansion of the universe

caused mainly by variations in tempera- uct and is therefore expressed in mature


ture between day and night. Pressure messenger RNA, ribosomal RNA, or trans-
changes in newly exposed rocks may also fer RNA. In eukaryotes, exons are sepa-
cause large-scale exfoliation. rated by noncoding sequences called
*introns.
exhalation See expiration.
exitance Symbol M. The radiant or lumi- exonuclease An enzyme that catalyses
the cleavage of nucleotides from the end
nous ux emitted per unit area of a sur-
face. The radiant exitance (Me) is measured of a nucleic acid molecule. Compare endo-
in watts per square metre (W m2), while nuclease.
e
the luminous exitance (Mv) is measured in exopeptidase A protein-digesting en-
lumens per square metre (lm m2). Exi- zyme that cleaves amino acids from the
tance was formerly called emittance. ends of a polypeptide chain. *Carboxypep-
exocarp See pericarp. tidase, which breaks down proteins in the
small intestine, is an example of an exo-
exocrine gland A gland that discharges peptidase. Compare endopeptidase.
its secretion into a body cavity (such as
the gut) or onto the body surface. Exam- exoskeleton A rigid external covering
ples are the *sebaceous and *sweat for the body in certain animals, such as
glands, the *mammary glands, and part of the hard chitinous cuticle of arthropods.
the pancreas. Exocrine glands are formed An exoskeleton protects and supports the
in the embryo from the invagination of body and provides points of attachment
epithelial cells. Their secretions pass ini- for muscles. The cuticle of arthropods
tially into a cavity (an alveolus or acinus) must be shed at intervals to allow growth
and then out through a duct or duct net- to occur (see ecdysis). Other examples of
work, along which the secretion may be- exoskeletons are the shells of molluscs
come modied by exchange with the and the bony plates of tortoises and ar-
blood across the duct epithelium. madillos. Compare endoskeleton.

exocytosis The passage of material exosphere See earths atmosphere.


from the inside of the cell to the cell sur- exothermic Denoting a chemical reac-
face within membrane-bound vesicles. tion that releases heat into its surround-
The membranes of the vesicles fuse with ings. Compare endothermic.
the plasma membrane, releasing their
contents to the exterior. Exocytosis is exotic atom 1. An atom in which an
used both for the removal of waste ma- electron has been replaced by another
terial from the cell and for secretion; for negatively charged particle, such as a
example of mucus by *goblet cells. Com- muon or *meson. In this case the negative
pare endocytosis. particle eventually collides with the nu-
cleus with the emission of X-ray photons.
exodermis See hypodermis. 2. A system in which the nucleus of an
exoergic Denoting a nuclear process atom has been replaced by a positively
that gives out energy. Compare endoergic. charged meson. Such exotic atoms have
to be created artically and are unstable.
exogamy The fusion of reproductive
cells from distantly related or unrelated exotoxin See toxin.
organisms, i.e. *outbreeding. Compare en- expansion The writing of a function or
dogamy. quantity as a *series of terms. The series
exogenous Describing substances, stim- may be nite or innite. See binomial the-
uli, etc., that originate outside an organ- orem; taylor series.
ism. For example, vitamins that cannot be
expansion of the universe The hy-
synthesized by an animal are said to be
pothesis, based on the evidence of the
supplied exogenously in the diet. Compare
*redshift, that the distance between the
endogenous.
galaxies is continuously increasing. The
exon A nucleotide sequence in a gene original theory, which was proposed in
that codes for part or all of the gene prod- 1929 by Edwin Hubble, assumes that the
expansivity 308

galaxies are ying apart as a consequence for determining the structures of mol-
of the big bang with which the universe ecules, etc.
originated. Several variants have since
expiration (exhalation) The process by
been proposed. See also big-bang theory;
which gas is expelled from the lungs (see
hubble constant.
respiratory movement). In mammals,
expansivity (thermal expansion) 1. Lin- the volume of the thoracic cavity is re-
ear expansivity is the fractional increase duced by contraction of the internal
in length of a specimen of a solid, per unit *intercostal muscles and relaxation of the
e rise in temperature. If a specimen in-
creases in length from l1 to l2 when its
muscles of the diaphragm, assisted by up-
ward pressure of the abdominal organs.
temperature is raised , then the expan- As a result, pressure in the lungs exceeds
sivity () is given by: atmospheric pressure and gas ows out of
the lungs, allowing the pressures to equal-
l2 = l1(1 + ). ize. Compare inspiration.
This relationship assumes that is inde- expiratory centre See ventilation
pendent of temperature. This is not, in centre.
general, the case and a more accurate re-
lationship is: explantation The removal of cells, tis-
sues, or organs of animals and plants for
l2 = l1(1 + a + b2 + c3), observation of their growth and develop-
where a, b, and c are constants. ment in appropriate culture media. See
2. Supercial expansivity is the fractional also tissue culture; organ culture.
increase in area of a solid surface caused explosive A compound or mixture that,
by unit rise in temperature, i.e. when ignited or detonated, undergoes a
A2 = A1(1 + ), rapid violent chemical reaction that pro-
duces large amounts of gas and heat,
where is the supercial expansivity. To a accompanied by light, sound and a high-
good approximation = 2. pressure shock wave. Low explosives burn
3. Volume expansivity is the fractional in- comparatively slowly when ignited, and
crease in volume of a solid, liquid, or gas are employed as propellants in rearms
per unit rise in temperature, i.e. and guns; they are also used in blasting.
V2 = V1(1 + ), Examples include *gunpowder and vari-
ous smokeless propellants, such as *cor-
where is the cubic expansivity and = dite. High explosives decompose very
3. For liquids, the expansivity observed rapidly to produce an uncontrollable
directly is called the apparent expansivity blast. Examples of this type include *dy-
as the container will also have expanded namite, *nitroglycerine, and *trinitro-
with the rise in temperature. The abso- toluene (TNT); they are exploded using a
lute expansivity is the apparent expansiv- detonator. Other high-power explosives
ity plus the volume expansivity of the include pentaerythritol tetranitrate
container. For the expansion of gases, see (PETN) and ammonium nitride/fuel oil
charles law. mixture (ANFO). Cyclonite (RDX) is a mili-
experiment A process or trial designed tary high explosive; mixed with oils and
to test a scientic theory. waxes, it forms a plastic explosive (such as
Semtex). See also Chronology.
expert system A computer program
that, using stored data, can reach conclu- exponent A number or symbol that in-
sions in a particular eld of knowledge; in dicates the power to which another num-
effect, it makes decisions. Thus unlike a ber or expression is raised. For example,
(x + y)n indicates that the expression (x + y)
*database, it processes the data before
is raised to the nth power; n is the expo-
presenting the result. The user can usu-
nent. Any number or expression in which
ally question the systems reasoning,
the exponent is zero is equal to 1, i.e.
often via a language processor. Such sys-
x0 = 1.
tems are used for troubleshooting in in-
dustry, for diagnosing medical disorders, exponential A function that varies as
309 exponential growth

EXPLOSIVES

9001000 Gunpowder developed in China.


1242 English monk Roger Bacon (122092) describes the preparation of
gunpowder (using an anagram).
c.1250 German alchemist Berthold Schwarz claims to have reinvented
gunpowder.
1771 French chemist Pierre Woulfe discovers picric acid (originally used as a
e
yellow dye).
1807 Scottish cleric Alexander Forsyth (17671843) discovers mercury
fulminate.
1833 French chemist Henri Braconnot (17811855) nitrates starch, making a
highly flammable compound (crude nitrocellulose).
1838 French chemist Thophile Pelouze (180767) nitrates paper, making
crude nitrocellulose.
1845 German chemist Christian Schnbein (17991868) nitrates cotton,
making nitrocellulose.
1846 Italian chemist Ascania Sobrero (181288) discovers nitroglycerine.
1863 Swedish chemist J. Wilbrand discovers trinitrotoluene (TNT).
Swedish chemist Alfred Nobel (183396) invents a detonating cap
based on mercury fulminate.
1867 Alfred Nobel invents dynamite by mixing nitroglycerine and
kieselguhr.
1871 German chemist Hermann Sprengel shows that picric acid can be used
as an explosive.
1875 Alfred Nobel invents blasting gelatin (nitroglycerine mixed with
nitrocellulose).
1885 French chemist Eugne Turpin discovers ammonium picrate (Mlinite).
1888 Alfred Nobel invents a propellant from nitroglycerine and nitrocellulose
(Ballistite).
1889 British scientists Frederick Abel (18261902) and James Dewar invent a
propellant (Cordite) similar to Ballistite.
1891 German chemist Bernhard Tollens (18411918) discovers
pentaerythritol tetranitrate (PETN).
1899 Henning discovers cyclotrimethylenetrinitramine (RDX or cyclonite).
1905 US army officer B. W. Dunn (18601936) invents ammonium picrate
explosive (Dunnite).
1915 British scientists invent amatol (TNT + ammonium nitrate).
1955 US scientists develop ammonium nitratefuel oil mixtures (ANFO) as
industrial explosives.

the power of another quantity. If y = ax, y ex = 1 + x + x2/2! + x3/3! + + xn/n! +


varies exponentially with x. The function
ex, also written as exp(x), is called the ex- exponential growth A form of *popu-
ponential function (see e). It is equal to the lation growth in which the rate of growth
sum of the exponential series, i.e. is related to the number of individuals
exposure meter 310

5000 extensive variable A quantity in a


*macroscopic system that is proportional
to the size of the system. Examples of ex-
tensive variables include the volume,
world population in millions

4000
mass, and total energy. If an extensive
variable is divided by an arbitrary exten-
3000 sive variable, such as the volume, an *in-
tensive variable results. A macroscopic
system can be described by one extensive
e 2000 variable and a set of intensive variables.
extensometer Any device for meas-
1000
uring the extension of a specimen of a
material under longitudinal stress. A com-
mon method is to make the specimen
form part of a capacitor, the capacitance
year 1800 1850 1900 1950 2000 of which will change with a change in the
specimens dimensions.
Graph showing exponential growth of the human
population extensor Any muscle that causes a limb
to extend. See voluntary muscle. Compare
present. Increase is slow when numbers flexor.
are low but rises sharply as numbers in-
crease. If population number is plotted external conversion A process in
against time on a graph a characteristic which molecules in electronically excited
J-shaped curve results (see graph). In ani- states pass to a lower electronic state
mal and plant populations, such factors as (which is frequently the ground state) by
overcrowding, lack of nutrients, and dis- colliding with other molecules. In this
ease limit population increase beyond a process the electronic energy is eventually
certain point and the J-shaped exponen- converted into heat. Since this process in-
tial curve tails off giving an S-shaped (sig- volves collisions, the rate at which it oc-
moid) curve. curs depends on how frequently collisons
occur. As a result, this process occurs
exposure meter A photocell that oper- much faster in liquids than in gases. It is
ates a meter to indicate the correct expo- sometimes called collision quenching.
sure for a specied lm in photography. It
enables the correct shutter speed and exteroceptor Any *receptor that de-
aperture to be chosen for any photo- tects external stimuli. Examples of extero-
graphic circumstances. Some cameras ceptors are the thermoreceptors in the
have a built-in exposure meter that auto- skin, which monitor the temperature
matically sets the aperture according to of the external environment. Compare
the amount of light available and the cho- interoceptor.
sen shutter speed. extinction 1. (in biology) The irre-
extended ASCII A set of characters versible condition of a species or other
group of organisms of having no living
with *ASCII values between 128 and 255.
representatives in the wild, which follows
These characters may include special sym-
the death of the last surviving individual
bols, graphics characters, and accented
of that species or group. Extinction may
characters. The assignment of extended
occur on a local or global level; it can re-
ASCII characters is not standard. It de-
sult from various human activities, includ-
pends on the particular computer system
ing the destruction of habitats or the
and may also depend on the font being
overexploitation of species that are
used.
hunted or harvested as a resource. Species
extender An inert substance added to a at the top of a *food chain (e.g. large birds
product (paint, rubber, washing powder, of prey) will be more prone to extinction
etc.) to dilute it (for economy) or to mod- since they exist in relatively small num-
ify its physical properties. bers and will be affected by a deleterious
311 eyepiece

change at any of the levels in the food are the *chorion, *amnion, *allantois,
chain. See also mass extinction. 2. (in be- and *yolk sac.
haviour) The termination of a behaviour
extraordinary ray See double refrac-
pattern that is no longer appropriate. For
tion.
example, dogs can be conditioned to sali-
vate when they hear a bell ring in the extrapolation An *approximation tech-
absence of a food stimulus (see condition- nique for nding the value of a function
ing). However, if the bell continues to be or measurement beyond the values al-
rung in the absence of food the dogs will ready known. If the values f(x0), f(x1),,
gradually stop salivating on hearing the f(xn) of a function of a variable x are e
bell. 3. (in physics) A reduction in the in- known in the interval [x0,xn], the value of
tensity of radiation as a result of absorp- f(x) for a value of x outside the interval
tion or scattering as it passes through [x0,xn] can be found by extrapolation. The
matter. techniques used in extrapolation are usu-
ally not as good as those used in *interpo-
extinction coefcient A measure of lation.
the extent by which the intensity of a
beam of light is reduced by passing extremely high frequency (EHF) A
through a distance d of a solution having radio frequency between 30 000 mega-
a molar concentration c of the dissolved hertz and 300 gigahertz.
substance. If the intensity of the light is extremophile A bacterium that thrives
reduced from I1 to I2, the extinction under extreme conditions, e.g. at very
coefcient is [log(I1/I2)]/cd. high or very low temperatures, or in very
salty or acidic environments. For example,
extracellular Located or occurring out-
certain archaebacteria (see archaea),
side the cell. *Cuticularization is an exam-
termed hyperthermophiles, live in hot
ple of an extracellular process.
springs at temperatures near or even
extracellular matrix (ECM) The vis- above 100C. The enzymes of such organ-
cous watery uid that surrounds cells in isms exhibit great stability and have been
animal tissues. Secreted by the cells them- extracted for use in laboratory and com-
selves, it is the medium through which mercial processes.
they receive materials (e.g. nutrients, hor- extrinsic semiconductor See semicon-
mones) from elsewhere in the body and ductor.
via which they communicate with other
cells. The ECM is the environment in eye The organ of sight. The most primi-
which cells migrate during tissue develop- tive eyes are the *eyespots of some unicel-
ment and it contains constituents that lular organisms. More advanced eyes are
bind cells together to maintain tissue in- the *ocelli and *compound eyes of arthro-
tegrity. It consists of glycoproteins, *colla- pods (e.g. insects). The cephalopod mol-
gens, and other structural components. luscs (e.g. the octopus and squid) and
The ECM is especially prominent in con- vertebrates possess the most highly devel-
nective tissues, such as bone, cartilage, oped eyes (see illustration). These nor-
and adipose tissue, in which it is some- mally occur in pairs, are nearly spherical,
times called ground substance. and lled with uid. Light is refracted by
the *cornea through the pupil in the *iris
extraction 1. The process of obtaining a and onto the *lens, which focuses images
metal from its ore. 2. The separation of a onto the retina. These images are received
component from a mixture by selective by light-sensitive cells in the retina (see
solubility. See partition. cone; rod), which transmit impulses to
extraembryonic membranes (embry- the brain via the optic nerve.
onic membranes) The tissues produced by eyepiece (ocular) The lens or system of
an animal *embryo for protection and nu- lenses in an optical instrument that is
trition but otherwise taking no part in its nearest to the eye. It usually produces a
development. The four membranes, magnied image of the previous image
which are called fetal membranes in man, formed by the instrument.
eyespot 312

retina

sclerotic
ciliary body
aqueous humour choroid
pupil
vitreous
lens
e cornea
humour fovea

iris
conjunctiva
blind spot

optic nerve

Structure of the vertebrate eye

eyespot (stigma) 1. A structure found to nd which has priority and similarly to


in some free-swimming unicellular algae the pair C and D. If the two groups of
and in plant reproductive cells that con- highest priority are on the same side of
tains orange or red pigments (carotenoids) the bond then the isomer is designated Z
and is sensitive to light. It enables the cell (from German zusammen, together). If they
to move in relation to a light source (see are on opposite sides the isomer is desig-
phototaxis). 2. A spot of pigment found nated E (German entgegen, opposite). The
in some lower animals, e.g. jellysh.
letters are used in the names of com-
eye tooth A *canine tooth in the upper pounds; for example (E)-butenedioic acid
jaw. (fumaric acid) and (Z)-butenedioic acid
EZ convention A convention for the (maleic acid). In compounds containing
description of a molecule showing cis two (or more) double bonds numbers are
trans isomerism (see isomerism). In a mol- used to designate the bonds (e.g. (2E, 4Z)-
ecule ABC=CDE, where A, B, D, and E are 2,4-hexadienoic acid). The system is less
substituent groups, the sequence rule (see ambiguous than the cis/trans system of
cip system) is applied to the pair A and B describing isomers.

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