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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
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
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.
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
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.
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
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.
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:
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
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
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
blood vessel
yolk sac
e umbilical cord
amnion
embryo
chorion
vagina
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
Energy bands
energy flow 292
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
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
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
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.
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
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
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