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Alternate titles: Animalia

By Virginia C. MaioranaSee All Article History

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Key People:

Charles Elton Carl E. Akeley Spencer Fullerton Baird Karl P. Schmidt Ross Granville Harrison

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animal social behaviour reptile fish animal behaviour insect

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Top Questions
What is an animal?
What are the two major groups of animals?
When did animals first appear?
What are the basic functional systems of animals?
How are animals different from plants and fungi?
Summary

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freshwater jellyfish

animal, (kingdom Animalia), any of a group of multicellular eukaryotic organisms (i.e.,

as distinct from bacteria, their deoxyribonucleic acid, or DNA, is contained in a
membrane-bound nucleus). They are thought to have evolved independently from the
unicellular eukaryotes. Animals differ from members of the two other kingdoms of
multicellular eukaryotes, the plants (Plantae) and the fungi (Mycota), in fundamental
variations in morphology and physiology. This is largely because animals have
developed muscles and hence mobility, a characteristic that has stimulated the further
development of tissues and organ systems.

Animals dominate human conceptions of life on Earth not simply by their size,

abundance, and sheer diversity but also by their mobility, a trait that humans share.
So integral is movement to the conception of animals that sponges, which
lack muscle tissues, were long considered to be plants. Only after their small movements
were noticed in 1765 did the animal nature of sponges slowly come to be recognized.

In size animals are outdone on land by plants, among whose foliage they may often hide.
In contrast, the photosynthetic algae, which feed the open oceans, are usually too small
to be seen, but marine animals range to the size of whales. Diversity of form, in contrast
to size, only impinges peripherally on human awareness of life and thus is less noticed.
Nevertheless, animals represent three-quarters or more of the species on Earth, a
diversity that reflects the flexibility in feeding, defense, and reproduction
which mobility gives them. Animals follow virtually every known mode of living that has
been described for the creatures of Earth.

Animals move in pursuit of food, mates, or refuge from predators, and this movement
attracts attention and interest, particularly as it becomes apparent that the behaviour of
some creatures is not so very different from human behaviour. Other than out of simple
curiosity, humans study animals to learn about themselves, who are a very recent
product of the evolution of animals.
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The animal kingdom

Animals evolved from unicellular eukaryotes. The presence of a nuclear membrane in
eukaryotes permits separation of the two phases
of protein synthesis: transcription (copying) of deoxyribonucleic acid (DNA) in the
nucleus and translation (decoding) of the message into protein in the cytoplasm.
Compared to the structure of the bacterial cell, this gives greater control over which
proteins are produced. Such control permits specialization of cells, each with identical
DNA but with the ability to control finely which genes successfully send copies into the
cytoplasm. Tissues and organs can thus evolve. The semirigid cell walls found in plants
and fungi, which constrain the shape and hence the diversity of possible cell types, are
absent in animals. If they were present, nerve and muscle cells, the focal point of animal
mobility, would not be possible.

A definition of animals

A characteristic of members of the animal kingdom is the presence of muscles and the
mobility they afford. Mobility is an important influence on how an organism obtains
nutrients for growth and reproduction. Animals typically move, in one way or another,
to feed on other living organisms, but some consume dead organic matter or even
photosynthesize by housing symbiotic algae. The type of nutrition is not as decisive as
the type of mobility in distinguishing animals from the other two multicellular
kingdoms. Some plants and fungi prey on animals by using movements based on
changing turgor pressure in key cells, as compared with the myofilament-based mobility
seen in animals. Mobility requires the development of vastly more elaborate senses and
internal communication than are found in plants or fungi. It also requires a different
mode of growth: animals increase in size mostly by expanding all parts of the body,
whereas plants and fungi mostly extend their terminal edges.

All phyla of the animal kingdom, including sponges, possess collagen, a triple helix
of protein that binds cells into tissues. The walled cells of plants and fungi are held
together by other molecules, such as pectin. Because collagen is not found among
unicellular eukaryotes, even those forming colonies, it is one of the indications that
animals arose once from a common unicellular ancestor.

The muscles that distinguish animals from plants or fungi are specializations of the actin
and myosin microfilaments common to all eukaryotic cells. Ancestral sponges, in fact,
are in some ways not much more complex than aggregations of protozoans that feed in
much the same way. Although the sensory and nervous system of animals is also made
of modified cells of a type lacking in plants and fungi, the basic mechanism of
communication is but a specialization of a chemical system that is found in protists,
plants, and fungi. The lines that divide an evolutionary continuum are rarely sharp.

Mobility constrains an animal to maintain more or less the same shape throughout its
active life. With growth, each organ system tends to increase roughly proportionately. In
contrast, plants and fungi grow by extension of their outer surfaces, and thus their shape
is ever changing. This basic difference in growth patterns has some interesting
consequences. For example, animals can rarely sacrifice parts of their bodies to satisfy
the appetites of predators (tails and limbs are occasionally exceptions), whereas plants
and fungi do so almost universally.
History of classification
Except perhaps for the possession of collagen, the criteria used above to distinguish
animals from other forms of life are not absolute. The first catalogs of
animal diversity were based on overall form and similarity. Aristotle and other early
biologists regarded all organisms as part of a great chain, divisions of which were more
or less arbitrary. The 18th-century Swedish botanist Carolus Linnaeus divided all
animals into six classes: Mammalia, Aves, Amphibia (including
reptiles), Pisces, Insecta (Arthropoda), and Vermes (other invertebrates). In the early
1800s the French zoologist Georges Cuvier recognized that vertebrates were
substantially different from invertebrates, and he divided most animals on the basis of
form and function into four branches: vertebrates, arthropods (articulates), mollusks,
and radiates (animals with radial symmetry). Cuvier’s divisions formed the basis for all
subsequent classifications.

Just after Cuvier’s classification, the French naturalist Étienne Geoffroy Saint-

Hilaire outlined the importance of homologous structures. Homology is correspondence
between features caused by continuity of information. Thus, a bird’s wing is homologous
to a bat’s wing insofar as both are forelimbs, but they are not homologous as wings.
Homologous structures need not resemble each other; for example, the three bones in
the middle ear of humans are homologous to three bones in the jaw apparatus in fishes
because the genetic and developmental information controlling them has been
continuous through evolutionary change.

Before evolution was generally accepted, homologies among different animals, when

they were recognized at all, were regarded as aspects of God’s pattern. Evolution
provided a testable explanation for homologies. By carefully tracing selected
homologies, it has been possible to show that previously
proposed classifications established inappropriate relationships based solely on form or
function, or both; for example, the radial symmetry of starfishes is not homologous to
that of coelenterates (such as jellyfish).

Protozoans were once considered to be animals because they move and do not

photosynthesize. Closer study has shown, though, that their movement is by means of
nonmuscular structures (cilia, flagella, or pseudopods) and that photosynthesis in them
has often been lost and gained. Protozoans do not, therefore, form a natural group but
with algae form a eukaryotic kingdom separate from plants and animals, called Protista.

Like plants and animals, fungi arose from protists and are now accorded a kingdom of
their own.

Animal diversity

early sea animals

The diverse appearance of animals is mostly superficial; the bewildering variety of

known forms, some truly bizarre, can be assorted among a mere half-dozen basic body
plans. These plans are established during the embryonic stages of development and
limit the size and complexity of the animals. Symmetry, number and relative
development of tissue layers, presence and nature of body cavities, and several aspects
of early development define these fundamental modes of organization.
Parazoa: a cellular level of organization

Although the two phyla in this subkingdom, Porifera (sponges) and Placozoa, lack

clearly defined tissues and organs, their cells specialize and integrate their activities.
Their simplicity has been adaptive, and sponges have remained important in benthic
marine habitats since their origin. The sessile, filter-feeding way of life shown by
sponges has favoured a body plan of radial symmetry, although some members have
become asymmetrical. The shape of the creeping, flattened placozoans is irregular and
changeable.
Radiata: a tissue level of organization
Study a jellyfish's muscular contractions and learn how the carnivore uses its tentacles to catch prey

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The two coelenterate phyla (Cnidaria and Ctenophora) advanced in complexity beyond

the parazoans by developing incipient tissues—groups of cells that are integrally
coordinated in the performance of a certain function. For example, coelenterates have
well-defined nerve nets, and their contractile fibres, although only specialized parts of
more generalized cells, are organized into discrete muscle units. Because discrete cells of
different types do not carry out the internal functions of the animals, coelenterates are
considered to be organized at only a tissue level.

The integration of cells into tissues, particularly those of nerve and muscle, permits a
significantly larger individual body size than is possible with other modes of body
movement. Flagella and cilia become ineffective at rather small size, and amoeboid
movement is limited to the size a single cell can attain. Muscles contract by a cellular
mechanism basically like that used in amoeboid locomotion—interaction of actin and
myosin filaments. Through coordinated contraction of many cells, movement of large
individuals becomes possible.

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Coelenterates, like parazoans, have only two body layers, an inner endoderm primarily

for feeding and an outer ectoderm for protection. Between the endoderm and the
ectoderm of coelenterates is the mesoglea, a gelatinous mass that contains connective
fibres of collagen and usually some cells. Both layers contain muscle fibres and a two-
dimensional web of nerve cells at the base; the endoderm surrounds a central cavity,
which ranges from simple to complex in shape and serves as a gut, circulatory system,
and sometimes even a skeleton. The cavity is also used for gamete dispersal and waste
elimination.

Cleavage of a fertilized egg produces a hollow sphere of flagellated cells (the blastula).

Invagination of cells at one or both poles creates a mouthless, solid gastrula; the gastrula
is called the planula larva in species in which this stage of development is free-living.
The inner, endoderm cells subsequently differentiate to form the lining of the central
cavity. The mouth forms once the planula larva has settled. Although the details of early
development are different for parazoans and coelenterates, most share a stage in which
external flagellated cells invaginate to form the inner layer, which lines the cavity, of
these diploblastic (two-layered) animals. This is characteristic of invagination during
the development of all animals.

All coelenterates are more or less radially symmetrical. A radial form is equally
advantageous for filtering, predatory, or photosynthetic modes of feeding. Tentacles
around the circumference can intercept food in all directions.
Bilateria: an organ level of organization
All animals except those in the four phyla mentioned above have bilaterally symmetrical
ancestors and contain three body layers (triploblastic) with coalition of tissues into
organs. The body plans that are generally recognized are acoelomate, pseudocoelomate,
and coelomate.

Acoelomates have no internal fluid-filled body cavity (coelom). Pseudocoelomates have

a cavity between the inner (endoderm) and the middle (mesoderm) body layers.
Coelomates have a cavity within the mesoderm, which can show one of two types of
development: schizocoelous or enterocoelic. Most protostomes show schizocoelous
development, in which the mesoderm proliferates from a single cell and divides to form
a mass on each side of the body; the coelom arises from a split within each mass.
Deuterostomes show enterocoelic pouching, in which the endoderm evaginates and
pinches off discrete pouches, the cavities of which become the coelom and the wall the
mesoderm. The animals in these major divisions of the Bilateria differ in other
fundamental ways, which are detailed below.

Unlike sessile sponges or floating jellyfish, the Bilateria typically move actively in

pursuit of food, although many members have further evolved into sessile or radial
forms. Directed movement is most efficient if sensory organs are located at the head or
forward-moving end of the animal. Organs of locomotion are most efficiently arranged
along both sides, a fact that defines the bilateral symmetry; many internal organs are
not in fact paired, whereas muscle layers, limbs, and sensory organs almost invariably
are. The diffuse nerve net of coelenterates coalesces into definite tracts or bundles,
which run posteriorly from the anterior brain to innervate the structures of locomotion.
Acoelomates
Flatworms (phyla Platyhelminthes, Nemertea, and Mesozoa) lack a coelom, although
nemerteans have a fluid-filled cavity at their anterior, or head, end, which is used to
eject the proboscis rapidly. The lack of a fluid-filled cavity adjacent to the muscles
reduces the extent to which the muscles can contract and the force they exert (see
below Support and movement). Because most also lack a circulatory system, supplying
muscle tissues with fuel and oxygen can be no faster than the rate at which these
substances diffuse through solid tissue. Flatworms are thus constrained to be relatively
flat and comparatively small; parasitic worms, which do not locomote, can achieve
immense lengths (e.g., tapeworms), but they remain very thin. The larger of the free-
living flatworms have extensively divided guts, which reach to within a few cells of the
muscles, thus compensating for the lack of a circulatory system. Most flatworms have
but one opening to the gut. Nemerteans, in addition to a coelom-like housing for their
proboscis, have attained a one-way gut and a closed circulatory system. Both increase
their ability to move food and oxygen to all parts of the body. Flatworms are considered
to be the ancestors of all other Bilateria.
Pseudocoelomates, or aschelminths
The pseudocoelomates include the nematodes, rotifers, gastrotrichs, and introverts.
Some members of some other phyla are also, strictly speaking, pseudocoelomate. These
four phyla of tiny body size (many species no larger than the bigger protozoans) are
placed together in part because they lack mesoderm on the inner side of the body cavity.
Consequently, no tissue, muscular or connective, supports the gut within the coelomic
fluid. For tiny organisms, this is advantageous for conservation of tissue: there is no
reason to evolve or to maintain a tissue that is not functionally important. The
inconspicuousness of most of these phyla has led to a slow advancement in
understanding their phylogenetic position in the animal kingdom.

Coelomates

reef squid
The advantage of a true coelom is the ability of the inner mesenteric (mostly connective
tissue) layer to suspend the central gut in the middle of the animal. Otherwise, in those
animals with a body cavity used in locomotion, gravity would pull the gut down and
severely curtail body size. Coelomates have attained vastly larger body sizes than has
any other group of animals. Within the coelomates, the coelom has been of variable
significance to the form and diversity of the various phyla. For example, it is essential
for the burrowing abilities of annelids and related phyla. It has largely lost this
significance in the arthropods, however, which have transferred locomotion to limbs
supported by an exoskeleton rather than a coelomic hydroskeleton. Suspension is the
main function of the coelom in vertebrates, which achieve the largest body sizes among
animals by virtue of an endoskeleton that does not need to be shed during growth.

The protostome coelomates (acoelomates and pseudocoelomates are also protostomes)

include the mollusks, annelids, arthropods, pogonophorans, apometamerans,
tardigrades, onychophorans, phoronids, brachiopods, and bryozoans. Deuterostomes
include the chaetognaths, echinoderms, hemichordates, and chordates.

In early development protostome coelomates mostly differ

from deuterostome coelomates in the following ways: (1) The mouth of protostomes is
the blastopore, the original opening into the developing gut which is formed during the
invagination of cells during gastrulation; that of deuterostomes is a secondary opening,
with the blastopore becoming the anus. (2, 3) Early cleavage is typically spiral and
determinate in protostomes, which means that the dividing cells are oriented at an angle
to one another and that the ultimate fate of the cells is mostly determined from the
beginning. Deuterostomes, in contrast, show indeterminate, radial cleavage, with the
dividing cells becoming layered and the fate of early cells a product of where they are
positioned later in development. (4) Coelom formation is schizocoelous in most
protostomes, whereas enterocoelous development is typical of deuterostomes. (5) For
those with a larval stage, the characteristic larval forms also differ.
The two phyla that have clearly dominated both land and sea since nearly the beginning
of animal evolution are the arthropods and chordates, protostomous and
deuterostomous coelomates, respectively. A key to arthropod success has been the
differentiation of many serially repeated parts, in particular jointed appendages with a
rigid exoskeleton, to perform the varied functions necessary to maintain life. The
exoskeleton, however, sets a moderate upper limit to body size. In contrast, vertebrates
share all habitats with arthropods by virtue of the larger maximum size permitted by the
development of an internal rigid skeleton. More than does a coelom, the evolution of
rigid, jointed skeletons has allowed these two phyla to dominate most
animal communities.
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Social levels of organization

Large size is often competitively advantageous but unobtainable by many animals

because of constraints of basic body plan. Intrinsically small animals sometimes become
large in the same way that protozoans evolved into metazoans: they multiply the
number of individuals by asexual reproduction (thus maintaining the same genotype)
and remain attached, with the option that individuals can be modified during their
development for a specialized function. This type of asexual sociality forms the
colonoids of sponges, coelenterates, bryozoans, hemichordates, and tunicate chordates,
all of which were primitively small, sessile filter feeders. Staying together
after asexual budding of new individuals gave a competitive edge to monopolizing
available space. With slight modifications so that all individuals in the colony could
share equally in the gains, these larger entities had the energy reserves necessary to
outcompete smaller organisms for space. This type of sociality has evolved in ways that
complicate the definition of individuality. For instance, Portuguese men-of-war and
their kin (some hydrozoan coelenterates) look and act like single individuals, yet their
components develop as genetically identical units, each homologous to a
whole jellyfish or polyp. It is a question whether such an animal should be considered
one individual or many.

A different type of sociality emerged among mobile complex animals that can
individually attain large size. In fact, the largest known living animals, the whales and
elephants, comprise two of a very few mammalian orders that contain only social
species. The pattern of evolution on Earth has favoured sociality in the smallest and the
largest (mostly vertebrates) of animals, albeit for different reasons. The smallest seek
the advantages of being large, as protozoans did to form the first animals. The large
animals can communicate; they spread out to find food, which all can share, and they
protect one another. Among the social groups of large animals, only humans
have differentiated their functions to such an extent that their societies begin to behave
as individuals.

Insect societies show behaviours halfway between societies based on genetically

identical members and those created by genetically different individuals; such
properties largely reflect their intermediate degree of genetic relatedness. Insects are
more cooperative and show a greater degree of altruism than is true
of vertebrate societies.

Form and function

animal lifespans

To stay alive, grow, and reproduce, an animal must find food, water, and oxygen, and it
must eliminate the waste products of metabolism. The organ systems typical of all but
the simplest of animals range from those highly specialized for one function to those
participating in many. The more basic functional systems are treated below from a
broadly comparative basis.
Support and movement
A skeleton can support an animal, act as an antagonist to muscle contraction, or, most
commonly, do both. Because muscles can only contract, they require some other
structure to stretch them to their noncontracted (relaxed) state. Another set of muscles
or the skeleton itself can act as an antagonist to muscle contraction. Only elastic
skeletons can act without an antagonist; all antagonistic muscles act through a skeleton,
which can be either rigid, flexible, or hydrostatic.
Types of skeletons and their distribution
Hydrostatic skeletons are the most prevalent skeletal system used by animals for
movement and support. A minimal hydroskeleton resembles a closed container. The
walls are two layers of muscles (antagonists) oriented at right angles to one another; the
inside contains an incompressible fluid or gel. The contraction of one set of muscles
exerts a pressure on the fluid, which is forced to move at right angles to the squeezing
antagonist. The movement of the fluid stretches the other set of muscles, which can then
contract to stretch its antagonist back to its relaxed position. The net result is an
alternating change in the shape of the container. Locomotion as varied as
crawling, burrowing, somersaulting, looping, or even walking is possible when the
container has some means of traction against a substrate: the system extends forward
from the point of attachment, attaches at a more forward point, releases posteriorly, and
contracts forward. Hydroskeletons are also important in nonlocomotory muscular
systems, such as hearts or intestines, which move blood or food, respectively.
Contraction-relaxation cycles push in one direction only when the system has structures
that prevent backflow.

Hydroskeletons become less efficient when fluid is lost. The optimal volume of fluid for
a particular system must remain constant for effective contraction and expansion of
the antagonistic muscles. If too much fluid is lost, the animal becomes limp and neither
muscle can stretch; when too much fluid is gained, the animal becomes bloated and
neither muscle can contract. Those coelenterates that use a hydroskeleton regularly face
a loss of pressure because their skeleton is also their gut. Freshwater animals tend to
become bloated as water diffuses into their salty cells, but terrestrial animals with
hydroskeletons tend to become limp as they dry. Solutions to water loss tend to be
partial because impermeable barriers, such as a shell, tend not to be very flexible, thus
negating the use of a hydroskeleton for movement. Terrestrial animals with locomotory
hydroskeletons (e.g., snails and earthworms) are restricted in their activity to moist
conditions.

Partitioning a hydroskeleton into many small, separate, but coordinated

units facilitates locomotion. In an earthworm, for example, a front group of segments
narrows together, thereby elongating that part of the worm. If there were no partitions
between the segments, the fluid would flow farther back, providing little elongation.
Widened segments behind these initial segments anchor the worm, and its head moves
forward. The process then reverses in a wave, and the posterior end moves
forward. Metamerism, or the partitioning of the coelom, is thought to have evolved in
ancestral annelids to improve their ability as burrowers in the bottom mud of the ocean.
It undoubtedly explains the unrivaled success of this phylum among worms and helps to
explain the extraordinary success of one of its relatives, the arthropods, which remained
segmented even after the skeletal function of the coelom was lost.

Elastic skeletons do not change shape but simply bend when a muscle contracts. Muscle
relaxation results either from a muscle contracting in the opposite direction to
its antagonist or from the skeleton resuming its original position. The tentacles of many
hydrozoan coelenterates, the mesoglea of jellyfish, the hinge of clamshells, and the
notochord of chordates are examples. The high-pressured coelom contained in the rigid
but flexible cuticle of nematodes also functions like an elastic skeleton.

Rigid, jointed skeletons achieve movement through a lever system. The elements of the
skeleton are rigid segments attached together by flexible joints. Muscles span the joints
and attach at each end to different elements. The more stable attachment site of a
muscle is called the origin, the other the insertion. One muscle contracts and moves the
skeletal element on which it is inserted, and an antagonistic muscle contracts and moves
the skeletal element in the opposite direction. The biceps and triceps of the upper arm in
humans are such a set of antagonistic muscles that bend and straighten, respectively,
the lower arm. The control of movement can be quite precise with jointed skeletons.
Muscles can bend or rotate skeletal elements whose length, shape, and number
contribute to the resulting action. The dexterity of the hands is an example of the
complexity of controlled movements made possible by a jointed skeleton.

Important to the speed and force of a movement are the length of the skeletal element
and the size of the contracting muscle. Short limbs with thick muscles have more power
than long limbs with slender muscles, but the latter have more speed. Limbs thus reveal
a great deal about how an animal moves. Likewise, the relative massiveness of jaws
reflects the toughness of the food eaten.

Two animal phyla, Chordata (vertebrates only) and Arthropoda, exploit jointed

skeletons. Although the skeleton is internal in vertebrates and external in arthropods,
the principles of movement are the same. A jointed skeleton is ideal for moving on land
because adaptations for protection against dehydration (such as the cuticle) do not
interfere with the action of the skeletal system. Indeed, the arthropod cuticle serves
jointly a protective and a skeletal role. Moreover, the diverse range of precise
movements made possible by this skeleton facilitates all sorts of locomotory patterns:
swimming, digging, running, climbing, and flying. Jointed skeletons are also used
directly for feeding (jaws). Arthropod jaws are derived from legs, while vertebrate jaws
are derived from gill arches.
Translating movement into locomotion and feeding
Although all animals can move, not all locomote or displace the body over a distance.
Locomotion serves the animal in finding food and mates and in escaping predators or
unsuitable habitats. These functions of locomotion are typically correlated among
different animals, so that those using the same mechanism of locomotion usually also
feed, seek mates, and avoid danger in similar ways.

Some of the correlations between mode of locomotion and mode of feeding are
described here, but space precludes discussion of the rich diversity found among
animals past and present. The locomotory/feeding system of animals is the heart of
their adaptation to their physical and biotic environments. Locomotory strategies for
finding or gathering food include the following techniques.

Sitting still and waiting for food to arrive is particularly prevalent in aquatic habitats but
is not rare on land. Sessile animals tend to develop strong defenses that are sometimes
incompatible with effective locomotion. They rely on water or air currents or on the
locomotion of their potential prey to bring food within reach. Because food may come
from any direction, many sessile animals evolve radial symmetry. Settlement may be
permanent or temporary, but in all cases one stage of the life cycle is capable of moving
actively or passively from its place of origin. The choice of attachment site can also be
active or passive; passive choice is often associated with an ability to grow in such a way
as to maximize feeding efficiency. As with plants, passive settlers do well only with luck.
The retention of locomotory capabilities requires energy and nutrients that can
otherwise be diverted into growth or the production of offspring. Sessile feeders need to
move if feeding and resting sites differ. Sessile animals include filter feeders, predators,
and even photosynthesizers; the latter include corals that house symbiotic algae.
Internal parasites are usually sessile because they live within their lifetime food supply.
Mobile animals that pursue sedentary strategies for seeking prey include web-spinning
spiders (a terrestrial mode of filter feeding) or deep-sea fishes with morphological
adaptations that lure prey.

Burrowing animals typically eat the rich organic substrates they move through. Others
burrow for protection and either temporarily emerge and gather organic sediments at
the top of their burrows or pump water with potential food through the burrow. Instead
of digging or finding burrows, some animals move into the centre of sponges, where
they find protection and a renewing source of food.

Active movement in search of food requires energy, but this expenditure is more than
made up for by an ability to seek out areas of concentrated food. This method of feeding
applies to burrowing animals that eat the substrate through which they move, as well as
to animals that move over solid surfaces, swim, or fly. Actively moving animals can feed
on organisms that do not move, a rich variety coating virtually the entire solid surface
of Earth, from the depths of the oceans to the peaks of many mountains. The main
problem with this most productive avenue of food gathering is protection. Shells and
poisons are the major types of defenses, although innovative detoxification metabolism
and jaws of various kinds breach the defenses in part. This is an escalating battle in
which the defenses, as well as the weapons to penetrate them are continually improving.
Nudibranchs, shell-less marine snails, incorporate the defensive stinging cells of prey
cnidarians into their own skin. Poisonous plants are eaten by specialized insects that
avoid or detoxify the poison. In fresh water, for reasons not known, the arms race has
not proceeded as far as in the sea.

Cooperation of individuals enables social animals to obtain food in novel ways.

Uncannily like humans, some ants farm and herd other organisms for food. For
example, some cultivate a fungus on leaves they cannot directly digest, while others herd
aphids from which they milk nectar (actually the phloem sap of plants). Some ants even
raid the nests of other species and make slaves of them. Another form of cooperation is
the mutualism between species that trade advantage for advantage. Some fishes feed on
parasites on the surfaces of other fishes, which benefits all but the parasites. In many
animals, including termites and ruminants, microorganisms thrive in the gut and digest
cellulose for them.

The nervous system
Coherent movement results only when the muscles receive a sensible pattern of
activating signals (for example, antagonists must not be activated to contract
simultaneously). Animals use specialized cells called neurons to coordinate their
muscular activity; nerves are bundles of neurons or parts thereof. Neurons
communicate between cells by chemical messengers, but within a single cell (often
extremely long) they can send high-speed signals through a wave of ionic polarization
(analogous to an electric current) along their membranes, a property inherent in all cells
but developed for speed in nerve cells by special modifications.

A system of communication requires three parts: a collector of outside information, an

integrator to evaluate that information and decide upon its relevance, and a transmitter
to convey the decision to the motor unit. In animals, sensory nerves and organs such as
eyes collect the information; associative nerves usually concentrated into a
brain integrate, evaluate, and decide its relevance; and effector or motor nerves convey
decisions to the muscles or elsewhere. Although all three parts of the nervous system
have kept pace with increases in the size and complexity of animals, the simplest
systems found among animals (those of parazoans and coelenterates) are nevertheless
capable of intricate feats of coordination. All ends of a coelenterate bipolar neuron can
both receive and transmit an impulse, whereas the unipolar neurons of more derived
animals receive only at one end (dendrite) and transmit at the other (axon). A neuron
can have multiple dendrites and axons.

The earliest animals were probably radial in design, so that bipolar neurons arranged in
a netlike pattern made sense. In such a design, a stimulus impinging at any point on the
body can travel everywhere to alert a simple array of myofilaments to contract
simultaneously. In the case of directed locomotion and relevant sensory input received
at the head end of a bilateral animal, unidirectional transmission of nerve impulses to
muscles becomes the only way to communicate effectively. The location of the brain in
the head also reflects efficiency and the speed of receipt of information, because this
position minimizes the distance between sensory and associative neurons as well as
concentrates these two functions in a small, protected part of the body. In most animals
nerve cells cannot be replaced if lost, although axons can be. Nerve cells tend to be
concentrated centrally in ganglia or nerve cords, with long axons extending peripherally.
Although certain animals may lose tails or limbs to predators or in accidents and then
regenerate them, loss or damage to the central nervous system means death or paralysis.

The nervous system uses the transmission properties of neurons to communicate.

Within a neuron, propagation of an impulse by an ion wave can be extremely rapid, but
the wave can pass along the length of only one cell’s membrane. To pass to the next cell
at a synapse, where an axon meets a dendrite, a chemical transmitter is required. This
molecule diffuses to the dendrites of a connecting neuron, where it initiates an ionic
wave that propagates along the length of the cell’s membrane. Although chemical
transmission is considerably slower than the ionic wave, it is more flexible. For example,
learning involves in part increasing the sensitivity of a particular nerve pathway to a
stimulus. The sensitivity of a synapse can be altered by increasing the amount of
transmitter released from the axon per impulse received, increasing the number of
receptors in the dendrite, or changing the sensitivity of the receptors. Bridging the
synapse directly by the formation of membrane-bound gap junctions, which
connect adjacent cells, enables an impulse to pass unimpeded to a connecting cell. The
increase in speed of transmission provided by a gap junction, however, is offset by a loss
in flexibility; gap junctions essentially create a single neuron from several. The same
result can be achieved more effectively by lengthening the axons or dendrites, making
some nerve cells metres in length. Situations arise where gap junctions become
desirable, however. Gap junctions are found in vertebrate cardiac and smooth muscles,
both of which transmit impulses along their cells to others. This ability makes these
muscles somewhat independent of nervous-system control. A body can thus be kept
partly functioning for some time without the activity of a brain.

Nerve impulses travel faster along axons of greater diameter or along those with good
insulation against ion leakage (except at spaced nodes required for recharging).
Vertebrates use their unique myelinated axons to increase the transmission rate of nerve
impulses, whereas invertebrates are limited to using axons of greater diameter. As a
result, vertebrates can concentrate more small neurons into a body of a particular size,
with the potential for greater complexity of behaviour.

Memory is still a poorly understood aspect of the nervous system. As in learning, both
short- and long-term memories seem to involve alterations in the ease with which
subsequent impulses travel a particular pathway after it has been used. Transfer of
memory through direct ingestion of the brain has not been confirmed experimentally.
Although the underlying mechanisms are only dimly understood, it is known that there
is a correlation between learning and memory capacity. The capacities for both increase
with the number of associative neurons and the number of branches or interconnections
formed. Since learning is a process of associating incoming cues with appropriate motor
or internal response, greater memory capacity of a brain gives a more rapid learning
process. Memory of inappropriate responses to an incoming set of cues can be used
without motor repeat.

The degree to which the neurons of a brain develop interconnections is correlated with
the complexity of its environs while growing. Consequently, a brain with fewer neurons
but with more interconnections can be more “intelligent” than one with more neurons.
Basic, repeated behaviours are inherited or learned by the development of fixed
pathways by which an environmental signal reaches the motor nerves rapidly with little
or no variation (reflex arcs). Nonreflex behaviour requires a decision to be made in the
brain, with the resulting pathway to the motor nerves becoming more fixed (habitual) as
one particular decision seems always to be correct. Reflexes are faster than decisions,
but their relative adaptiveness depends on context. Animals vary in the degree to which
they use reflexes or make decisions, patterns that are strongly correlated to brain size.
Habitual actions are perhaps the most prevalent response, a compromise between the
speed of a response and its appropriateness to context.
The senses
Appropriate behaviour relies on receiving adequate information from
the environment to alert an animal to the presence of food, mates, or danger. Although
sensory nerves carry this information to the brain, they do not always directly perceive
the external world. Other modified cells intervene to convert light waves into vision,
pressure waves in air or water into sound, chemicals into smell or taste, and simple
contact into touch. Some animals have other senses, as for electric or magnetic fields.

In vision, for example, a photosensitive molecule changes shape and thereby sets off a
chain of reactions that ultimately depolarize the dendrite of a sensory nerve. The
associative neurons in the brain interpret the pattern of incoming impulses into a
composite picture. What is “seen” may not entirely map what is really there: a great deal
of filtering occurs, with editing by the brain to eliminate less important details so that
only the most important are perceived. The accuracy of what is seen increases with brain
size and the complexity of the visual gathering system, or eyes. Animal eyes range from
being able to discern only the presence or absence of light to being able to see objects in
vivid colour and great detail. Some animals see in ranges beyond unaided human vision.
Pollinating insects in particular discern the colour of flowers differently than do
humans; the ultraviolet reflection patterns of flowers do not always coincide with their
coloured ones. Bees and birds perceive polarized light and can orient themselves by it.
Some animals perceive long wavelengths, which are associated with heat (infrared), and
can locate the presence of warm-blooded prey by such a mechanism.

Chemoreceptors are usually little-modified sensory neurons, except for the taste

receptors of vertebrates, which are frequently replaced cells in synaptic contact with
permanent sensory neurons. Chemoreception is based on the recognition of molecules
at receptor sites, lipid-protein complexes that are liberally scattered on the dendrites of
a sensory neuron. When the receptor recognizes one particular molecule by shape and
sometimes chemical composition, it fires an impulse. The pattern of firings set off in the
receptors of a certain molecule provides the information that the brain interprets as an
odour or a taste. The details of how animals smell and taste are not as well understood
as are the other senses. In many animals, chemoreceptors are not concentrated into
obvious organs as they are in vertebrates, making even their location difficult to discern.
Most animals possess some sort of chemoreception, and in many the sense is a major
part of the animal’s perception of its environment, far more so than it is for humans.

Sounds are waves of molecular disturbance that move through air, water, or solids, and
their perception by animals simply uses sensitive mechanoreceptors. (Loud sounds can
also be felt by the general touch receptors of the body and thereby influence its sense of
well-being.) Sound receptors are sensitive hair cells or membranes that depolarize a
sensory neuron when bent by the passage of a sound wave. Direct deformation of the
dendritic membrane or release of transmitters by the hair cells fire the sensory neurons.
Aside from a few insects, only vertebrates have organs with which to hear. Fishes and
aquatic amphibians use a lateral-line system, and other vertebrates use ears; both
organs use hair cells as phonoreceptors. Sound waves directly stimulate the hair cells of
lateral-line systems, while sound waves only indirectly stimulate the hair cells of ears
through an amplifying system of membranes and bones, which reaches a peak of
complexity in mammals. Some animals (e.g., most bats and whales and even whirligig
beetles) use sound to “see” by echolocation. Sound is the preferred medium of
communication between animals that hear. It can be used over longer distances than
vision, and it can be used when vision is not possible. The signals decay more rapidly
than do those of odours, and therefore the information can be more precise.
Mechanoreceptors also respond to touch, pressure, stretching, and gravity. They are
located all over the body and enable an animal to monitor its state at any moment. Much
of this monitoring is subconscious but necessary for normal functioning.
Mechanoreceptors are often just sensory nerves, but other cells may be involved. Unlike
other senses, that of touch is found in all animals, even sponges, where it reflects a
general cellular trait of eukaryotes.

Hormones
Hormones are the chemical integrators of a multicellular existence, coordinating
activities from daily maintenance to reproduction and development. The
neurotransmitters released by axons are one class of chemical communicators that act
on an adjacent cell, usually a muscle cell or another neuron. Hormones are a mostly
distinct class of chemical communicators secreted by nerves, ordinary tissue, or special
glands; they act on cells far removed from the site of their release. They can be proteins,
single polypeptides, amines, or steroids or other lipids. Hormones travel to their place of
action via the circulatory system and then match their particular configuration with a
specific receptor molecule attached to a cell membrane or, more usually, located within
the cell.

The nervous system coordinates the more rapid activities of animal life, such as

movement, while the hormones integrate everything else. Only the larger, more complex
animals, such as vertebrates and some arthropods, have special endocrine glands to
produce hormones; other animals use nerve cells or tissues such as the gonads.
Endocrine glands are another example of a partitioning of functions into separate
organs, a system that increases efficiency but that requires a relatively large size to
maintain. Greater specialization is also associated with greater difficulties in
regenerating lost parts or preventing breakdowns in functions.

Although the list of hormones found in the mammalian body may seem large, the
numbers are surprisingly low for the variety of functions they influence. Which of the
multiple functions any one hormone regulates depends on the specificity of the
receptors on or within cells. Because all hormones bathe all cells as a result of their
transport by the circulatory system, it is more efficient to have a general messenger
transported to a cell, where it elicits only one of many possible outcomes. As in the
nervous system, the specificity of response lies in the organ that responds and not with
the messenger that merely commands action.

Chemicals that allow communication among individuals are called pheromones. Sexual
attractants are the most common, but there are many other kinds.
Digestion
In contrast to plants, the essential nutrients that animals require to sustain life and to
reproduce come packaged with their source of energy—the flesh or organic remains of
other organisms. More complex animals tend to shorten and even eliminate
many synthetic pathways, because most of the essential building blocks of their own
complex molecules are present in their food. Reducing synthetic flexibility,
however, inhibits a radical alteration in diet. The digestive and synthetic chemistry of
animals strongly reflects their diets; some of this design may be altered with diet, and
some may not. No matter how many leafy vegetables humans consume, for example, the
cellulose remains undigested because appropriate microorganisms are not present in
the digestive tract and they cannot be obtained at will. Consequently, essential nutrients
are species-specific and tend to include only molecules adequately available in the usual
diet.

The structure of a digestive system reflects its typical diet. Its purpose is to process food
only to the point at which it can be transported to other cells for use as either fuel or
structural material. In the simplest animals, such as sponges or some coelenterates,
digestion is entirely intracellular, and some of the products of digestion are transported
to nondigestive cells. As animals began to catch larger types of food, more of the
digestive process had to be handled extracellularly. At the simplest level, seen in
coelenterates or flatworms, large food items are held in an internal cavity (the gut) or
even externally where certain cells release digestive enzymes. The food is broken down
only to the stage at which it can be ingested by cells, which finish the process
intracellularly. In more complex animals extracellular digestion accounts for virtually all
breakdown of food before the products are transported to nondigestive cells.

Chemical digestion, whether intracellular or extracellular, is a relatively slow way

to decompose a large item. Thus, animals begin to break it apart mechanically before
exposing it to digestive enzymes. Teeth, the molluscan radula, and muscular gizzards are
organs that speed up the digestive process by macerating food into finer particles.

Very early in their evolution animals acquired a one-way gut (gastrointestinal system),

with the mouth typically armed with the macerating equipment and the terminal stretch
sometimes specialized to retrieve excess water or other nutrients. Often a single passage
through the digestive system leaves a great deal of useful material unclaimed. Because
food moves along at a characteristic rate, which is sometimes influenced by how much is
coming in, not all can be fully digested. Some animals regularly eat their feces to retrieve
nutrients that may have escaped during first passage. If not recycled by their owners,
feces are consumed by a diverse set of organisms.

A common specialization of the gut is the stomach or crop—a highly extensible part of

the digestive tract that is used to hold a large amount of food and partially digest it
before it enters the intestines, where most of the chemical breakdown and absorption of
nutrients occurs. Most animals eat intermittently; the less often they eat, the larger the
relative stomach size. Internalizing as much food as possible when it is available
prevents potential food from being taken by a stronger competitor or enables a feeder to
retreat to safety while digesting its meal. Ceca and second stomachs provide symbiotic
microorganisms with a safe area within the gut to digest cellulose. Excess
microorganisms mixed in with the partly digestible wastes contribute a steady protein-
rich fare to the host in exchange for an optimal place to consume cellulose.

Stomachs predominate as a gut specialization because they allow animals to keep food
from competitors or other dangers, but a few animals have developed ingenious
methods of digesting their food before ingesting it. Humans are latecomers to this
practice and have not yet carried it very far. Starfish exploit secondary
radial symmetry and tube feet to open bivalved mollusks only enough to inject their
stomachs, digest their meal within the protected shell, absorb the products, and leave
the wastes behind. Spiders immobilize prey by silk wrappings and venoms, inject
digestive enzymes, and drink the brew. Some primitive animals, like placozoans and
certain flatworms, simply hunch over their prey as they digest it externally, a practice
that leaves them vulnerable to other predators.

Animals use surfaces in many ways but no more strikingly than in the gut. Nutrients
enter the body proper through the surface membrane of the gut; the larger the animal,
the larger this surface area must be. The gut is probably the system that best reflects an
animal’s ecology. The simplest guts, found in animals from sponges to flatworms, simply
branch like trees as the animal increases in size; the gut itself reaches all parts of the
body to within the distance of a few cells and thus can serve for nutrient transport. As
muscle masses become more prominent, the gut is squeezed into a more compact form.
The gut compensates for this lack of space by internalizing its foldings. For example, the
lining of the mammalian small intestine, the major site of digestion and absorption, is
not only folded but each cell also has numerous outpocketings (microvilli), which
increase the surface area 25-fold. Mammals and birds that primarily eat plants have
longer intestines than those that favour meat. Warm-blooded animals, which maintain
constant internal temperatures, require a great deal more energy than cold-blooded
ones and thus tend to concentrate more surface area into a gut. Although they are not
efficient energy users, it is to their advantage to obtain more usable energy even if
efficiency is lost in the process.