ANIMAL EVOLUTION – INVERTEBRATES

I. Medicines from the Sea

Animal life began in the sea, and the oceans remain the greatest repository of animal diversity. In the oceans,
as on land, the majority of animals are invertebrates, meaning they do not have a backbone. Only about 5
percent of animals have a backbone and thus are considered vertebrates. Invertebrates evolved long before
vertebrates, and they remain the most diverse and numerous animals. The longevity and diversity of
invertebrate lineages attests to how well they have adapted to their environment. Many marine invertebrates
produce secondary metabolites that help them survive. These compounds may protect an animal from
predators, help it fend off pathogens, or assist in the capture of prey. Some such compounds also have effects
in the human body, and so can be useful as medicines. Consider the venom that some fish-eating cone snails
inject into their prey to subdue it. The snail’s venom anesthetizes and paralyzes a fish, thus preventing it from
struggling with and possibly harming the snail as it is captured and consumed. Human nerves and fish nerves
use the same chemical communication signals, so compounds used by a cone snail to drug fish can also affect
the function of the human nervous system. A person stung by a fish-eating cone snail may become numb at
the site of injection, suffer from temporary paralysis, or even die. A synthetic version of one peptide in cone
snail venom is now used as a pain reliever. The drug, ziconotide (Prialt), is injected to suppress severe pain
that cannot be controlled by other means. Other peptides isolated from cone snail venom are being tested as
treatments for epilepsy, diabetes, or cancer. Compounds derived from other marine invertebrates are also
already in use. AZT (azidothymidine), the first drug successfully used to treat AIDS, is a synthetic derivative of a
molecule first discovered in a sponge. Other compounds from sponges can be used to treat infections caused
by herpesviruses. Gorgonians, which are relatives of sea anemones, are the source of a variety of anti-
inflammatory compounds, one of which is used in an
“anti-aging” face cream.

II. Animal Traits and Body Plans

1. What is an Animal?
Animals are multicelled heterotrophs that take food
into their body, where they digest it and absorb the
released nutrients. An animal body consists of a few to hundreds of types of unwalled cells. These cells
become specialized as an animal develops from an embryo (an individual in the earliest stage of development)
to an adult. Most animals reproduce sexually, some reproduce asexually, and some do both. All are motile
(move from place to place) during part or all of their life.

2. Variation in Animal Body Plans

All animals are multicellular and constitute the clade Metazoa. The earliest animals were probably
aggregations of cells, and this level of organization persists in
sponges. However, most modern animals have cells organized as
tissues. Tissue organization begins in animal embryos. Embryos
of jellies and other cnidarians have two tissue layers: an outer
ectoderm and an inner endoderm. In other modern animals,
embryonic cells typically rearrange themselves to form a middle
tissue layer called mesoderm. Evolution of a three-layer embryo
allowed an important increase in structural complexity. Most
internal organs in animals develop from embryonic mesoderm.
Animals with the simplest structural organization, such as
sponges, are asymmetrical; their body cannot be divided into
two halves that are mirror images. Jellies, sea anemones, and
other cnidarians have radial symmetry: Body parts are repeated
around a central axis, like spokes of a wheel. Radial animals
usually attach to an underwater surface or drift along. A radial
body plan allows them to capture food that can arrive from any
direction. Animals with a three-layer body plan typically have
bilateral symmetry: The body’s left and right halves are mirror
images. Such lineages typically undergo cephalization, an
evolutionary process whereby many nerve cells and sensory structures become concentrated at the front of
the body. These structures help the animal find food or avoid threats as it moves headfirst through its
environment. The two lineages of bilateral animals are defined in part by developmental differences. In
protostomes, the first opening that appears on an embryo becomes a mouth. Proto– means first and stoma
means opening. Most bilateral invertebrates are protostomes. In deuterostomes, the mouth develops from
the second embryonic opening. Deuterostomes include some invertebrates, and all vertebrates. Animals also
differ in how they digest food. In sponges, digestion is intracellular. Cnidarians and flatworms digest food
inside a saclike gut called a gastrovascular cavity. Food enters this cavity through the same opening that expels
wastes. The cavity also functions in gas exchange. Most bilateral animals have a tubular gut, or complete
digestive tract, with an opening at either end. Parts of the tube specialize in taking in food, digesting it,
absorbing nutrients, or compacting waste. Unlike a gastrovascular cavity, a tubular gut carries out all of these
tasks simultaneously. A mass of tissues and organs surrounds a flatworm’s gut. Most other animals have a
“tube within a tube” body plan. Their gut runs through a fluid-filled body cavity. In roundworms, this cavity is
partially lined with tissue derived from mesoderm, and is called a pseudocoelom. More typically, bilateral
animals have a coelom, a body cavity fully lined with tissue derived from mesoderm. Sheets of tissue called
mesentery suspend the gut in the center of a coelom. Coelomic fluid cushions the gut and keeps it from being
distorted by body movements. The fluid also helps distribute material through the body and in some animals,
it plays a role in locomotion.

Like plants, all animals are multicelled. Like fungi, they are consumers. Unlike either group, animals can move.

3. Colonial Origins
According to the colonial theory of animal origins, the first animals evolved from a type of colonial protist. At
first, all cells in the colony were similar. Each could reproduce and carry out all other essential tasks. Later,
mutations resulted in cells that were better at some tasks but did not carry out others. Perhaps these cells
captured food more efficiently, but did not reproduce. Colonies that had interdependent cells and a division of
labor were at a selective advantage. Over time, new specialized cell types evolved. Eventually this process
produced the first animal. What was the protist ancestral to animals like? Choanoflagellates, the modern
protists most closely related to animals, provide some clues. Choanoflagellates are flagellated cells that live
either as single cells or as a colony of genetically identical cells. In their structure, choanoflagellate cells closely
resemble some cells in the bodies of modern sponges.

4. Early Animals
Chemicals discovered in rocks found during an oil drilling
project in Oman provide the earliest evidence of animals.
The sedimentary rocks, which were laid down as seafloor
more than 635 million years ago, contain a biomarker
characteristic of sponges. A biomarker is a distinctive
molecule produced only by a particular type of cell or
lineage. Similarly aged rocks from Australia contain what
appear to be fossils of sponge bodies. A collection of 570-
million-year-old fossil organisms first discovered in Australia’s Ediacara Hills provides evidence of an early
animal diversification. These fossils, collectively known as Ediacarans, include a variety of soft-bodied
organisms. Many of these fossils appear to be marine invertebrates, and some are thought to be early
representatives of modern invertebrate groups. Others belong to groups that have no surviving members.

5. The Cambrian Explosion

Animals underwent a dramatic adaptive radiation during the Cambrian (542–488 million years ago). By the
end of this period, all major animal lineages were present in the seas. What caused this Cambrian explosion in
diversity? Environmental factors probably played a role. During this period, the global climate warmed and the
oxygen concentration rose in the seas. Both factors made the environment more hospitable to animal life.
With rare exceptions, animals require oxygen. In modern seas, highly oxygenated waters support a greater
species diversity than cooler ones. Also, during the Cambrian, the supercontinent Gondwana underwent a
dramatic rotation. Movement of this landmass could have isolated populations, thus increasing the likelihood
that allopatric speciation events would occur. Biological factors could also have encouraged diversification.
After predatory animals arose, evolution of novel prey defenses such as protective hard parts would have
been favored. Duplications and divergence of homeotic genes would have facilitated diversification. Changes
in these genes have dramatic effects on body plans. Some mutations produced adaptive traits that better
allowed animals to more easily escape predators or to survive in novel habitats.
Fossils and gene comparisons among modern species provide insights into how animals arose and diversified.

III. Sponges
Sponges (phylum Porifera) are aquatic animals that do
not have tissues or organs. Most of the more than
5,000 species live in tropical seas. Some are as small as
a fingertip, whereas others stand meters tall. An
asymmetrical vaselike or columnar shape is most
common, but some grow as a thin crust. An adult
sponge is a sessile animal, meaning it lives attached to
a surface. The body has many pores. Flat,
nonflagellated cells cover the body’s outer surface,
flagellated collar cells line its inner surface, and a
jellylike extracellular matrix fills the space in between.
Amoeba-like sponge cells live in the matrix. Many
sponges also have cells that secrete fibrous proteins or
glassy silica spikes. These materials structurally support
the body, and help fend off predators. Some protein-
rich sponges are harvested from the sea, dried,
cleaned, and bleached. Their rubbery protein remains
are used for bathing and cleaning. The typical sponge is
a suspension feeder, meaning it filters its food from
the surrounding water. The movement of flagella on
collar cells draws food-laden water through pores in a
sponge’s body wall. The collar cells filter food from the
water and engulf it by phagocytosis. Digestion is intracellular. The amoeba-like cells in the matrix receive
breakdown products of digestion from collar cells, and distribute these nutrients to other cells in the sponge
body. Most sponges are hermaphrodites, meaning each individual can produce both eggs and sperm.
Typically, a sponge releases sperm into the water but holds on to its eggs. Fertilization produces a zygote that
develops into a ciliated larva. A larva (plural, larvae) is a young, sexually immature animal with a body form
that differs from that of an adult. Sponge larvae swim briefly, then settle and develop into adults. Many
sponges can also reproduce asexually when small buds or fragments break away and grow into new sponges.
Some freshwater sponges can survive dry conditions by producing gemmules, which are tiny clumps of resting
cells encased in a hardened coat. Gemmules are dispersed by the wind. Those that land in a hospitable habitat
become active and grow into new sponges.

IV. Body Plans

Cnidarians (phylum Cnidaria) include 10,000 species of radially symmetrical animals such as corals, sea
anemones, and jellies (also called jellyfishes). Nearly all are marine. There are two cnidarian body plans—
medusa and polyp. In both, a tentacle-ringed orifice opens onto a gastrovascular cavity that functions in both
 digestion and gas exchange. A medusa (plural, medusae) is shaped like
a bell or umbrella, with a mouth on the ventral (lower) surface. Most
swim or drift about. A polyp is tubular, and one end usually attaches to
a surface. Cnidarians have an embryo with two tissue layers. Their outer
epidermis develops from embryonic ectoderm, and the inner
gastrodermis from endoderm. Mesoglea, a jellylike secreted matrix, fills
the space between these tissue layers. Cnidarians are predators that
use their tentacles to sting and capture prey. The name Cnidaria is from
cnidos, the Greek word for nettle, a kind of stinging plant. The tentacles
have cells called cnidocytes with specialized organelles (nematocysts)
that act like a jack-in-the-box. When something brushes against the cell,
a coiled thread pops out and entangles prey or sticks a venomous barb into it. The captured prey is pushed
through the mouth, into the saclike gastrovascular cavity.
Gland cells of the gastrodermis secrete enzymes that
digest the prey, then digestive remains are expelled
through the mouth. Cnidarians are brainless, but
interconnecting nerve cells extend through their tissues,
forming a nerve net. Body parts move when nerve cells
signal contractile cells. In a manner analogous to
squeezing a waterfilled balloon, the contractions
redistribute mesoglea or water trapped in the
gastrovascular cavity. A fluid-filled cavity or cellular mass on which contractile cells exert force is a hydrostatic
skeleton.

V. Diversity and Life Cycles

There are four cnidarian classes: hydrozoans, anthozoans, cubozoans, and scyphozoans. Some marine
hydrozoans have a life cycle that includes polyp, medusa, and larval stages. A cnidarian larva, called a planula,
is ciliated and bilateral. Hydra, another hydrozoan, is a freshwater predatory polyp about 20 millimeters (3/4
inch) tall. There is no medusa stage, and reproduction usually occurs asexually by budding. Anthozoans such as
corals and sea anemones also do not have a medusa stage. Gametes form on polyps. Coral reefs consist of
colonies of polyps that enclose themselves in a framework of secreted calcium carbonate. Photosynthetic
dinoflagellates live in the polyp’s tissues and provide it with sugars. Cubozoans and scyphozoans have a
medusa-dominated life cycle. Cubozoans, commonly called box jellies, are active swimmers with structurally
complex eyes, complete with a lens. Box jellies that live in the Indo-Pacific Ocean produce a powerful venom,
and their sting can be deadly to humans. Scyphozoans include most jellies that commonly wash up on
beaches. Some are harvested and dried for use as food, especially in Asia. The Portuguese man-of-war
(Physalia) is a large colonial scyphozoan that consists of many individuals.