Animals are a major group of multicellular, eukaryotic
organisms of the kingdom Animalia or Metazoa. Their body plan eventually
becomes fixed as they develop, although some undergo a process of metamorphosis
later on in their life. Most animals are motile, meaning they can move
spontaneously and independently. All animals are also heterotrophs, meaning
they must ingest other organisms or their products for sustenance.
Most known animal phyla appeared in the fossil record as
marine species during the Cambrian explosion, about 542 million years ago.
Etymology
The word "animal" comes from the Latin word
animalis, meaning "having breath".[1] In everyday colloquial usage,
the word often refers to non-human members of kingdom Animalia. Sometimes, only
closer relatives of humans such as mammals and other vertebrates are meant in
colloquial use.[2] The biological definition of the word refers to all members
of the kingdom Animalia, encompassing creatures as diverse as sponges,
jellyfish, insects and humans.[3]
Characteristics
Animals have several characteristics that set them apart
from other living things. Animals are eukaryotic and multicellular,[4] which
separates them from bacteria and most protists. They are heterotrophic,[5]
generally digesting food in an internal chamber, which separates them from
plants and algae.[6] They are also distinguished from plants, algae, and fungi
by lacking rigid cell walls.[7] All animals are motile,[8] if only at certain
life stages. In most animals, embryos pass through a blastula stage,[9] which
is a characteristic exclusive to animals.
Structure
With a few exceptions, most notably the sponges (Phylum
Porifera) and Placozoa, animals have bodies differentiated into separate
tissues. These include muscles, which are able to contract and control
locomotion, and nerve tissues, which send and process signals. Typically, there
is also an internal digestive chamber, with one or two openings.[10] Animals
with this sort of organization are called metazoans, or eumetazoans when the
former is used for animals in general.[11]
All animals have eukaryotic cells, surrounded by a
characteristic extracellular matrix composed of collagen and elastic
glycoproteins.[12] This may be calcified to form structures like shells, bones,
and spicules.[13] During development, it forms a relatively flexible framework[14]
upon which cells can move about and be reorganized, making complex structures
possible. In contrast, other multicellular organisms, like plants and fungi,
have cells held in place by cell walls, and so develop by progressive
growth.[10] Also, unique to animal cells are the following intercellular
junctions: tight junctions, gap junctions, and desmosomes.[15]
Reproduction and development
See also: Sexual reproduction#Animals and Asexual
reproduction#Examples in animals
A newt lung cell stained with fluorescent dyes undergoing
the early anaphase stage of mitosis
Nearly all animals undergo some form of sexual
reproduction.[16] They have a few specialized reproductive cells, which undergo
meiosis to produce smaller, motile spermatozoa or larger, non-motile ova.[17]
These fuse to form zygotes, which develop into new individuals.[18]
Many animals are also capable of asexual reproduction.[19]
This may take place through parthenogenesis, where fertile eggs are produced
without mating, budding, or fragmentation.[20]
A zygote initially develops into a hollow sphere, called a
blastula,[21] which undergoes rearrangement and differentiation. In sponges,
blastula larvae swim to a new location and develop into a new sponge.[22] In
most other groups, the blastula undergoes more complicated rearrangement.[23]
It first invaginates to form a gastrula with a digestive chamber, and two
separate germ layers — an external ectoderm and an internal endoderm.[24] In
most cases, a mesoderm also develops between them.[25] These germ layers then
differentiate to form tissues and organs.[26]
Food and energy sourcing
Main article: Animal nutrition
All animals are heterotrophs, meaning that they feed
directly or indirectly on other living things.[27] They are often further
subdivided into groups such as carnivores, herbivores, omnivores, and
parasites.[28]
Predation is a biological interaction where a predator (a
heterotroph that is hunting) feeds on its prey (the organism that is
attacked).[29] Predators may or may not kill their prey prior to feeding on
them, but the act of predation always results in the death of the prey.[30] The
other main category of consumption is detritivory, the consumption of dead
organic matter.[31] It can at times be difficult to separate the two feeding behaviours,
for example, where parasitic species prey on a host organism and then lay their
eggs on it for their offspring to feed on its decaying corpse. Selective
pressures imposed on one another has led to an evolutionary arms race between
prey and predator, resulting in various antipredator adaptations.[32]
Most animals indirectly use the energy of sunlight by eating
plants or plant-eating animals. Most plants use light to convert inorganic
molecules in their environment into carbohydrates, fats, proteins and other
biomolecules, characteristically containing reduced carbon in the form of
carbon-hydrogen bonds. Starting with carbon dioxide (CO2) and water (H2O),
photosynthesis converts the energy of sunlight into chemical energy in the form
of simple sugars (e.g., glucose), with the release of molecular oxygen. These
sugars are then used as the building blocks for plant growth, including the
production of other biomolecules.[10] When an animal eats plants (or eats other
animals which have eaten plants), the reduced carbon compounds in the food
become a source of energy and building materials for the animal.[33] They are
either used directly to help the animal grow, or broken down, releasing stored
solar energy, and giving the animal the energy required for motion.[34][35]
Animals living close to hydrothermal vents and cold seeps on
the ocean floor are not dependent on the energy of sunlight.[36] Instead
chemosynthetic archaea and bacteria form the base of the food chain.[37]
Origin and fossil record
Further information: Urmetazoan
Dunkleosteus was a gigantic, 10-metre-long (33 ft)
prehistoric fish.[38]
Vernanimalcula guizhouena is a fossil believed by some to
represent the earliest known member of the Bilateria.
Animals are generally considered to have evolved from a
flagellated eukaryote.[39] Their closest known living relatives are the
choanoflagellates, collared flagellates that have a morphology similar to the
choanocytes of certain sponges.[40] Molecular studies place animals in a
supergroup called the opisthokonts, which also include the choanoflagellates,
fungi and a few small parasitic protists.[41] The name comes from the posterior
location of the flagellum in motile cells, such as most animal spermatozoa,
whereas other eukaryotes tend to have anterior flagella.[42]
The first fossils that might represent animals appear in the
Trezona Formation at Trezona Bore, West Central Flinders, South Australia.[43]
These fossils are interpreted as being early sponges. They were found in
665-million-year-old rock.[43]
The next oldest possible animal fossils are found towards
the end of the Precambrian, around 610 million years ago, and are known as the
Ediacaran or Vendian biota.[44] These are difficult to relate to later fossils,
however. Some may represent precursors of modern phyla, but they may be
separate groups, and it is possible they are not really animals at all.[45]
Aside from them, most known animal phyla make a more or less
simultaneous appearance during the Cambrian period, about 542 million years
ago.[46] It is still disputed whether this event, called the Cambrian
explosion, represents a rapid divergence between different groups or a change
in conditions that made fossilization possible.
Some paleontologists suggest that animals appeared much
earlier than the Cambrian explosion, possibly as early as 1 billion years
ago.[47] Trace fossils such as tracks and burrows found in the Tonian era
indicate the presence of triploblastic worms, like metazoans, roughly as large
(about 5 mm wide) and complex as earthworms.[48] During the beginning of the
Tonian period around 1 billion years ago, there was a decrease in Stromatolite
diversity, which may indicate the appearance of grazing animals, since
stromatolite diversity increased when grazing animals went extinct at the End
Permian and End Ordovician extinction events, and decreased shortly after the
grazer populations recovered. However the discovery that tracks very similar to
these early trace fossils are produced today by the giant single-celled protist
Gromia sphaerica casts doubt on their interpretation as evidence of early
animal evolution.[49][50]
It has been estimated that 99.9% of animals that have ever
existed are extinct.[51]
Groups of animals
The relative number of species contributed to the total by
each phylum of animals.
Porifera, Radiata and basal Bilateria
Phylogenetic analysis suggests that the Porifera and
Ctenophora diverged before a clade that gave rise to the Bilateria, Cnidaria
and Placozoa.[52] Another study based on the presence/absence of introns
suggests that Cnidaria, Porifera and Placozoa may be a sister group of
Bilateria and Ctenophora.[53]
Orange elephant ear sponge, Agelas clathrodes, in
foreground. Two corals in the background: a sea fan, Iciligorgia schrammi, and
a sea rod, Plexaurella nutans.
The sponges (Porifera) were long thought to have diverged
from other animals early.[54] They lack the complex organization found in most
other phyla.[55] Their cells are differentiated, but in most cases not
organized into distinct tissues.[56] Sponges typically feed by drawing in water
through pores.[57] Archaeocyatha, which have fused skeletons, may represent
sponges or a separate phylum.[58] However, a phylogenomic study in 2008 of 150
genes in 29 animals across 21 phyla revealed that it is the Ctenophora or comb
jellies which are the basal lineage of animals, at least among those 21 phyla.
The authors speculate that sponges—or at least those lines of sponges they
investigated—are not so primitive, but may instead be secondarily
simplified.[59]
Among the other phyla, the Ctenophora and the Cnidaria,
which includes sea anemones, corals, and jellyfish, are radially symmetric and
have digestive chambers with a single opening, which serves as both the mouth
and the anus.[60] Both have distinct tissues, but they are not organized into
organs.[61] There are only two main germ layers, the ectoderm and endoderm,
with only scattered cells between them. As such, these animals are sometimes
called diploblastic.[62] The tiny placozoans are similar, but they do not have
a permanent digestive chamber.
The remaining animals form a monophyletic group called the
Bilateria. For the most part, they are bilaterally symmetric, and often have a
specialized head with feeding and sensory organs. The body is triploblastic,
i.e. all three germ layers are well-developed, and tissues form distinct
organs. The digestive chamber has two openings, a mouth and an anus, and there
is also an internal body cavity called a coelom or pseudocoelom. There are
exceptions to each of these characteristics, however — for instance adult
echinoderms are radially symmetric, and certain parasitic worms have extremely
simplified body structures.
Genetic studies have considerably changed our understanding
of the relationships within the Bilateria. Most appear to belong to two major
lineages: the deuterostomes and the protostomes, the latter of which includes
the Ecdysozoa, Platyzoa, and Lophotrochozoa. In addition, there are a few small
groups of bilaterians with relatively similar structure that appear to have
diverged before these major groups. These include the Acoelomorpha, Rhombozoa,
and Orthonectida. The Myxozoa, single-celled parasites that were originally
considered Protozoa, are now believed to have developed from the Medusozoa as
well.
Deuterostomes
Superb Fairy-wren, Malurus cyaneus
Deuterostomes differ from the other Bilateria, called
protostomes, in several ways. In both cases there is a complete digestive
tract. However, in protostomes, the first opening of the gut to appear in
embryological development (the archenteron) develops into the mouth, with the
anus forming secondarily. In deuterostomes the anus forms first, with the mouth
developing secondarily.[63] In most protostomes, cells simply fill in the
interior of the gastrula to form the mesoderm, called schizocoelous development,
but in deuterostomes, it forms through invagination of the endoderm, called
enterocoelic pouching.[64] Deuterostome embryos undergo radial cleavage during
cell division, while protostomes undergo spiral cleavage.[65]
All this suggests the deuterostomes and protostomes are
separate, monophyletic lineages. The main phyla of deuterostomes are the
Echinodermata and Chordata.[66] The former are radially symmetric and
exclusively marine, such as starfish, sea urchins, and sea cucumbers.[67] The
latter are dominated by the vertebrates, animals with backbones.[68] These
include fish, amphibians, reptiles, birds, and mammals.[69]
In addition to these, the deuterostomes also include the
Hemichordata, or acorn worms.[70] Although they are not especially prominent today,
the important fossil graptolites may belong to this group.[71]
The Chaetognatha or arrow worms may also be deuterostomes,
but more recent studies suggest protostome affinities.
Ecdysozoa
Yellow-winged darter, Sympetrum flaveolum
The Ecdysozoa are protostomes, named after the common trait
of growth by moulting or ecdysis.[72] The largest animal phylum belongs here,
the Arthropoda, including insects, spiders, crabs, and their kin. All these
organisms have a body divided into repeating segments, typically with paired
appendages. Two smaller phyla, the Onychophora and Tardigrada, are close
relatives of the arthropods and share these traits.
The ecdysozoans also include the Nematoda or roundworms,
perhaps the second largest animal phylum. Roundworms are typically microscopic,
and occur in nearly every environment where there is water.[73] A number are
important parasites.[74] Smaller phyla related to them are the Nematomorpha or
horsehair worms, and the Kinorhyncha, Priapulida, and Loricifera. These groups have
a reduced coelom, called a pseudocoelom.
The remaining two groups of protostomes are sometimes
grouped together as the Spiralia, since in both embryos develop with spiral
cleavage.
Platyzoa
Pseudobiceros bedfordi, (Bedford's flatworm)
The Platyzoa include the phylum Platyhelminthes, the
flatworms.[75] These were originally considered some of the most primitive
Bilateria, but it now appears they developed from more complex ancestors.[76] A
number of parasites are included in this group, such as the flukes and
tapeworms.[75] Flatworms are acoelomates, lacking a body cavity, as are their
closest relatives, the microscopic Gastrotricha.[77]
The other platyzoan phyla are mostly microscopic and
pseudocoelomate. The most prominent are the Rotifera or rotifers, which are
common in aqueous environments. They also include the Acanthocephala or
spiny-headed worms, the Gnathostomulida, Micrognathozoa, and possibly the
Cycliophora.[78] These groups share the presence of complex jaws, from which
they are called the Gnathifera.
Lophotrochozoa
Roman snail, Helix pomatia
The Lophotrochozoa include two of the most successful animal
phyla, the Mollusca and Annelida.[79][80] The former, which is the
second-largest animal phylum by number of described species, includes animals
such as snails, clams, and squids, and the latter comprises the segmented
worms, such as earthworms and leeches. These two groups have long been
considered close relatives because of the common presence of trochophore
larvae, but the annelids were considered closer to the arthropods because they
are both segmented.[81] Now, this is generally considered convergent evolution,
owing to many morphological and genetic differences between the two phyla.[82]
The Lophotrochozoa also include the Nemertea or ribbon
worms, the Sipuncula, and several phyla that have a ring of ciliated tentacles
around the mouth, called a lophophore.[83] These were traditionally grouped
together as the lophophorates.[84] but it now appears that the lophophorate
group may be paraphyletic,[85] with some closer to the nemerteans and some to
the molluscs and annelids.[86][87] They include the Brachiopoda or lamp shells,
which are prominent in the fossil record, the Entoprocta, the Phoronida, and
possibly the Bryozoa or moss animals.[88]
Model organisms
Main articles: Model organism and Animal testing
Because of the great diversity found in animals, it is more
economical for scientists to study a small number of chosen species so that
connections can be drawn from their work and conclusions extrapolated about how
animals function in general. Because they are easy to keep and breed, the fruit
fly Drosophila melanogaster and the nematode Caenorhabditis elegans have long
been the most intensively studied metazoan model organisms, and were among the
first life-forms to be genetically sequenced. This was facilitated by the
severely reduced state of their genomes, but as many genes, introns, and
linkages lost, these ecdysozoans can teach us little about the origins of
animals in general. The extent of this type of evolution within the superphylum
will be revealed by the crustacean, annelid, and molluscan genome projects
currently in progress. Analysis of the starlet sea anemone genome has
emphasised the importance of sponges, placozoans, and choanoflagellates, also
being sequenced, in explaining the arrival of 1500 ancestral genes unique to
the Eumetazoa.[89]
An analysis of the homoscleromorph sponge Oscarella carmela
also suggests that the last common ancestor of sponges and the eumetazoan
animals was more complex than previously assumed.[90]
Other model organisms belonging to the animal kingdom
include the house mouse (Mus musculus) and zebrafish (Danio rerio).
Carolus Linnaeus, known as the father of modern taxonomy
History of classification
Aristotle divided the living world between animals and
plants, and this was followed by Carolus Linnaeus (Carl von Linné), in the
first hierarchical classification.[91] Since then biologists have begun
emphasizing evolutionary relationships, and so these groups have been
restricted somewhat. For instance, microscopic protozoa were originally
considered animals because they move, but are now treated separately.
In Linnaeus's original scheme, the animals were one of three
kingdoms, divided into the classes of Vermes, Insecta, Pisces, Amphibia, Aves,
and Mammalia. Since then the last four have all been subsumed into a single
phylum, the Chordata, whereas the various other forms have been separated out.
The above lists represent our current understanding of the group, though there
is some variation from source to source.
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