Superphylum Ecdysozoa: Nematodes and Tardigrades

Phylum Nematoda

The Nematoda, like other members of the superphylum Ecdysozoa, are triploblastic and possess an embryonic mesoderm that is sandwiched between the ectoderm and endoderm. They are also bilaterally symmetrical, meaning that a longitudinal section will divide them into right and left sides that are superficially symmetrical. In contrast with flatworms, nematodes are pseudocoelomates and show a tubular morphology and circular cross-section. Nematodes include both free-living and parasitic forms.

In 1914, N.A. Cobb said, “In short, if all the matter in the universe except the nematodes were swept away, our world would still be dimly recognizable, and if, as disembodied spirits, we could then investigate it, we should find its mountains, hills, vales, rivers, lakes and oceans represented by a thin film of nematodes...” To paraphrase Cobb, nematodes are so abundant that if all the non-nematode matter of the biosphere were removed, there would still remain a shadow of the former world outlined by nematodes!Stoll, N. R., “This wormy world. 1947,” Journal of Parasitology 85(3) (1999): 392-396. The phylum Nematoda includes more than 28,000 species with an estimated 16,000 being parasitic in nature. However, nematologists believe there may be over one million unclassified species.

The name Nematoda is derived from the Greek word “Nemos,” which means “thread,” and includes all true roundworms. Nematodes are present in all habitats, typically with each species occurring in great abundance. The free-living nematode, Caenorhabditis elegans, has been extensively used as a model system for many different avenues of biological inquiry in laboratories all over the world.


The cylindrical body form of the nematodes is seen in Figure. These animals have a complete digestive system with a distinct mouth and anus, whereas only one opening is present in the digestive tract of flatworms. The mouth opens into a muscular pharynx and intestine, which leads to a rectum and anal opening at the posterior end. The epidermis can be either a single layer of cells or a syncytium—a multinucleated tissue that in this case is formed by the fusion of many single cells. The cuticle of nematodes is rich in collagen and a polymer called chitin, which forms a protective armor outside the epidermis. The cuticle extends into both ends of the digestive tract, the pharynx, and rectum. In the head, an anterior mouth opening is composed of three (or six) “lips” as well as teeth derived from the cuticle (in some species). Some nematodes may present other modifications of the cuticle such as rings, head shields, or warts. These external rings, however, do not reflect true internal body segmentation, which as we have seen is a hallmark of phylum Annelida. The attachment of the muscles of nematodes differs from that of most animals: they have a longitudinal layer only, and their direct attachment to the dorsal and ventral nerve cords creates a strong muscular contraction that results in a whiplike, almost spastic, body movement.

Photo a shows a worm-shaped nematode next to a capsule-shaped nematode egg. The illustration in part b shows a cross-section of a nematode, which has a mouth at one end and an anus at the other. The mouth connects to a pharynx, then to an intestine. A dorsal nerve runs along the top of the animal and joins ring-like head ganglia at the front end. Testes run alongside the intestine toward the back of the animal.
Nematode morphology. Scanning electron micrograph shows (a) the soybean cyst nematode (Heterodera glycines) and a nematode egg. (b) A schematic representation shows the anatomy of a typical nematode. (credit a: modification of work by USDA ARS; scale-bar data from Matt Russell)

Excretory System

In nematodes, specialized excretory systems are not well developed. Nitrogenous wastes, largely in the form of ammonia, are released directly across the body wall. In some nematodes, osmoregulation and salt balance are performed by simple excretory cells or glands that may be connected to paired canals that release wastes through an anterior pore. In marine nematodes, the excretory cells are called renette cells, which are unique to nematodes.

Nervous system

Most nematodes have four longitudinal nerve cords that run along the length of the body in dorsal, ventral, and lateral positions. The ventral nerve cord is better developed than the dorsal and lateral cords. Nonetheless, all nerve cords fuse at the anterior end, to form a pharyngeal nerve ring around the pharynx, which acts as the head ganglion or the “brain” of the roundworm. A similar fusion forms a posterior ganglion at the tail. In C. elegans, the nervous system accounts for nearly one-third of the total number of cells in the animal!


Nematodes employ a variety of reproductive strategies ranging from monoecious to dioecious to parthenogenetic, depending upon the species. C. elegans is a mostly monoecious species with both self-fertilizing hermaphrodites and some males. In the hermaphrodites, ova and sperm develop at different times in the same gonad. Ova are contained in a uterus and amoeboid sperm are contained in a spermatheca ("sperm receptacle"). The uterus has an external opening known as the vulva. The female genital pore is near the middle of the body, whereas the male genital pore is nearer to the tip. In anatomical males, specialized structures called copulatory spicules at the tail of the male keep him in place and open the vulva of the female into which the amoeboid sperm travel into the spermatheca.

Fertilization is internal, and embryonic development starts very soon after fertilization. The embryo is released from the vulva during the gastrulation stage. The embryonic development stage lasts for 14 hours; development then continues through four successive larval stages with molting and ecdysis taking place between each stage—L1, L2, L3, and L4—ultimately leading to the development of a young adult worm. Adverse environmental conditions such as overcrowding or lack of food can result in the formation of an intermediate larval stage known as the dauer larva. An unusual feature of some nematodes is eutely: the body of a given species contains a specific number of cells as the consequence of a rigid developmental pathway.

Everyday Connection

C. elegans: The Model System for Linking Developmental Studies with Genetics If biologists wanted to research how nicotine dependence develops in the body, how lipids are regulated, or observe the attractant or repellant properties of certain odors, they would clearly need to design three very different experiments. However, they might only need one subject of study: Caenorhabditis elegans. The nematode C. elegans was brought into the focus of mainstream biological research by Dr. Sydney Brenner. Since 1963, Dr. Brenner and scientists worldwide have used this animal as a model system to study many different physiological and developmental mechanisms.

C. elegans is a free-living nematode found in soil. Only about a millimeter long, it can be cultured on agar plates (10,000 worms/plate!), feeding on the common intestinal bacterium Escherichia coli (another long-term resident of biological laboratories worldwide), and therefore can be readily grown and maintained in a laboratory. The biggest asset of this nematode is its transparency, which helps researchers to observe and monitor changes within the animal with ease. It is also a simple organism with about 1,000 cells and a genome of only 20,000 genes. Its chromosomes are organized into five pairs of autosomes plus a pair of sex chromosomes, making it an ideal candidate with which to study genetics. Since every cell can be visualized and identified, this organism is useful for studying cellular phenomena like cell-to-cell interactions, cell-fate determinations, cell division, apoptosis (cell death), and intracellular transport.

Another tremendous asset is the short life cycle of this worm (Figure). It takes only three days to achieve the “egg to adult to daughter egg”; therefore, the developmental consequences of genetic changes can be quickly identified. The total life span of C. elegans is two to three weeks; hence, age-related phenomena are also easy to observe. There are two sexes in this species: hermaphrodites (XX) and males (XO). However, anatomical males are relatively infrequently obtained from matings between hermaphrodites, since their XO chromosome composition requires meiotic nondisjunction when both parents are XX. Another feature that makes C. elegans an excellent experimental model is that the position and number of the 959 cells present in adult hermaphrodites of this organism is constant. This feature is extremely significant when studying cell differentiation, cell-to-cell communication, and apoptosis. Lastly, C. elegans is also amenable to genetic manipulations using molecular methods, rounding off its usefulness as a model system.

Biologists worldwide have created information banks and groups dedicated to research using C. elegans. Their findings have led, for example, to better understandings of cell communication during development, neuronal signaling, and insight into lipid regulation (which is important in addressing health issues like the development of obesity and diabetes). In recent years, studies have enlightened the medical community with a better understanding of polycystic kidney disease. This simple organism has led biologists to complex and significant findings, growing the field of science in ways that touch the everyday world.

Photo a shows transparent worm about a millimeter in length. Illustration B shows the life cycle of C. elegans, which begins when the egg hatches, releasing a L1 juvenile. After 12 hours the L1 juvenile transforms into an L2 juvenile. After 7 hours the L2 juvenile transforms into an L3 juvenile. After another 7 hours the L3 juvenile transforms into an L4 juvenile. After 14 hours the L4 juvenile transforms into an adult. The hermaphroditic adult mates with another adult to produce fertilized eggs which hatch, completing the cycle.
Caenorhabditis elegans. (a) This light micrograph shows the bodies of a group of roundworms. These hermaphrodites consist of exactly 959 cells. (b) The life cycle of C. elegans has four juvenile stages (L1 through L4) and an adult stage. Under ideal conditions, the nematode spends a set amount of time at each juvenile stage, but under stressful conditions, it may enter a dauer state that does not age significantly and is somewhat analogous to the diapausing state of some insects. (credit a: modification of work by “snickclunk”/Flickr: credit b: modification of work by NIDDK, NIH; scale-bar data from Matt Russell)

Parasitic Nematodes

A number of common parasitic nematodes serve as prime examples of parasitism (endoparasitism). These economically and medically important animals exhibit complex life cycles that often involve multiple hosts, and they can have significant medical and veterinary impacts. Here is a partial list of nasty nematodes: Humans may become infected by Dracunculus medinensis, known as guinea worms, when they drink unfiltered water containing copepods (Figure), an intermediate crustacean host. Hookworms, such as Ancylostoma and Necator, infest the intestines and feed on the blood of mammals, especially of dogs, cats, and humans. Trichina worms (Trichinella) are the causal organism of trichinosis in humans, often resulting from the consumption of undercooked pork; Trichinella can infect other mammalian hosts as well. Ascaris, a large intestinal roundworm, steals nutrition from its human host and may create physical blockage of the intestines. The filarial worms, such as Dirofilaria and Wuchereria, are commonly vectored by mosquitoes, which pass the infective agents among mammals through their blood-sucking activity. One species, Wuchereria bancrofti, infects the lymph nodes of over 120 million people worldwide, usually producing a non-lethal but deforming condition called elephantiasis. In this disease, parts of the body often swell to gigantic proportions due to obstruction of lymphatic drainage, inflammation of lymphatic tissues, and resulting edema. Dirofilaria immitis, a blood-infective parasite, is the notorious dog heartworm species.

 Part A shows a foot with a guinea worm extending from a blister. The end of the worm is wrapped around a stick. Part B shows the life cycle of the guinea worm, which begins when a person drinks unfiltered water containing copepods infected with guinea worm larvae. Larvae, which are released when the copepods die, penetrate the wall of the stomach and intestine. The worms mature and reproduce. Fertilized females migrate to the surface of the skin, where they discharge larvae into the water. Copepods consume the larvae. The copepods are consumed by humans, completing the cycle. About a year after infection, the female worm emerges from the skin.
Life cycle of the guinea worm. The guinea worm Dracunculus medinensis infects about 3.5 million people annually, mostly in Africa. (a) Here, the worm is wrapped around a stick so it can be slowly extracted. (b) Infection occurs when people consume water contaminated by infected copepods, but this can easily be prevented by simple filtration systems. (credit: modification of work by CDC)