Superphylum Lophotrochozoa: Flatworms, Rotifers, and Nemerteans

Phylum Platyhelminthes

The flatworms are acoelomate organisms that include many free-living and parasitic forms. The flatworms possess neither a lophophore nor trochophore larvae, although the larvae of one group of flatworms, the Polycladida (named after its many-branched digestive tract), are considered to be homologous to trochophore larvae. Spiral cleavage is also seen in the polycladids and other basal flatworm groups. The developmental pattern of some of the free-living forms is obscured by a phenomenon called "blastomere anarchy," in which a sort of temporary feeding larva forms, followed by a regrouping of cells within the embryo that gives rise to a second-stage embryo. However, both the monophyly of the flatworms and their placement in the Lophotrochozoa has been supported by molecular analyses.

The Platyhelminthes consist of two monophyletic lineages: the Catenulida and the Rhabditophora. The Catenulida, or "chain worms," is a small clade of just over 100 species. These worms typically reproduce asexually by budding. However, the offspring do not fully detach from the parents and the formation resembles a chain in appearance. All of the flatworms discussed here are part of the Rhabditophora ("rhabdite bearers"). Rhabdites are rodlike structures discharged in the mucus produced by some free-living flatworms; Eucoelmate protostomes are schizocoels, in which mesoderm-producing cells typically migrate into the blastocoel during gastrulation likely serve in both defense and to provide traction for ciliary gliding along the substrate. Unlike free-living flatworms, many species of trematodes and cestodes are parasitic, including important parasites of humans.

This image shows two flatworms with significant physical diversity.
Flatworms exhibit significant diversity. (a) A blue Pseudoceros flatworm (Pseudoceros bifurcus); (b) gold speckled flatworm (Thysanozoon nigropapillosum). (credit a: modification of work by Stephen Childs; b: modification of work by Pril Fish.)

Flatworms have three embryonic tissue layers that give rise to epidermal tissues (from ectoderm), the lining of the digestive system (from endoderm), and other internal tissues (from mesoderm). The epidermal tissue is a single layer of cells or a layer of fused cells (syncytium) that covers two layers of muscle, one circular and the other longitudinal. The mesodermal tissues include mesenchymal cells that contain collagen and support secretory cells that produce mucus and other materials at the surface. Because flatworms are acoelomates, the mesodermal layer forms a solid mass between the outer epidermal surface and the cavity of the digestive system.

Physiological Processes of Flatworms

The free-living species of flatworms are predators or scavengers. Parasitic forms feed by absorbing nutrients provided by their hosts. Most flatworms, such as the planarian shown in Figure, have a branching gastrovascular cavity rather than a complete digestive system. In such animals, the “mouth” is also used to expel waste materials from the digestive system, and thus also serves as an anus. (A few species may have a second anal pore or opening.) The gut may be a simple sac or highly branched. Digestion is primarily extracellular, with digested materials taken into the cells of the gut lining by phagocytosis. One parasitic group, the tapeworms (cestodes), lacks a digestive system altogether, and absorb digested food from the host.

Flatworms have an excretory system with a network of tubules attached to flame cells, whose cilia beat to direct waste fluids concentrated in the tubules out of the body through excretory pores. The system is responsible for the regulation of dissolved salts and the excretion of nitrogenous wastes. The nervous system consists of a pair of lateral nerve cords running the length of the body with transverse connections between them. Two large cerebral ganglia—concentrations of nerve cell bodies at the anterior end of the worm—are associated with photosensory and chemosensory cells. There is neither a circulatory nor a respiratory system, with gas and nutrient exchange dependent on diffusion and cell-to-cell junctions. This necessarily limits the thickness of the body in these organisms, constraining them to be “flat” worms. Most flatworm species are monoecious (both male and female reproductive organs are found in the same individual), and fertilization is typically internal. Asexual reproduction by fission is common in some groups.

Illustration shows the digestive, nervous and excretory systems in a flat, worm-like planaria. The digestive system starts at the ventral mouth opening in the middle of the animal, and then extends to the head through the middle of the body, and toward the along the sides of the body. Many lateral branches occur along the digestive system. The nervous system has 2 cerebral ganglia at the eyes in the head, and 2 ventral nerve cords with transverse connections along the length of the body to the tail. The excretory system is arranged in 2 long mesh-like structures down each side of the body.
Planaria, a free-living flatworm. The planarian is a flatworm that has a gastrovascular cavity with one opening that serves as both mouth and anus. The excretory system is made up of flame cells and tubules connected to excretory pores on both sides of the body. The nervous system is composed of two interconnected nerve cords running the length of the body, with cerebral ganglia and eyespots at the anterior end.

Diversity of Flatworms

The flatworms have been traditionally divided into four classes: Turbellaria, Monogenea, Trematoda, and Cestoda (Figure). However, the relationships among members of these classes has recently been reassessed, with the turbellarians in particular now viewed as paraphyletic, since its descendants may also include members of the other three classes. Members of the clade or class Rhabditophora are now dispersed among multiple orders of Platyhelminthes, the most familiar of these being the Polycladida, which contains the large marine flatworms; the Tricladida (which includes Dugesia [“planaria”] and Planaria and its relatives); and the major parasitic orders: Monogenea (fish ectoparasites), Trematoda (flukes), and Cestoda (tapeworms), which together form a monophyletic clade.

Photo A shows a Bedford’s flatworm from the class Turbellaria. The worm has the appearance of a fringed ribbon, black with pink stripes, swimming above the sand. Photo B shows a Dactylogyrus from the class Monogenea. The worm’s body is a long, thin translucent oval with bulges at one end that give the appearance of a head. Three dark spots appear in the head, and four more dark spots three-quarters appear of the way down the body. Anchors that enable the worm to latch onto gills are located near these spots. Photo C shows a foot- shaped brown worm. Photo D shows a long, thin ribbon-like white worm.
Traditional flatworm classes. Phylum Platyhelminthes was previously divided into four classes. (a) Class Turbellaria includes the free-living polycladid Bedford’s flatworm (Pseudobiceros bedfordi), which is about 8 to 10 cm in length. (b) The parasitic class Monogenea includes Dactylogyrus spp, commonly called gill flukes, which are about 0.2 mm in length and have two anchors, indicated by arrows used to attach the parasite on to the gills of host fish. (c) The class Trematoda includes Fascioloides magna (right) and Fasciola hepatica (two specimens on left, also known as the common liver fluke). (d) Class Cestoda includes tapeworms such as this Taenia saginata, infects both cattle and humans, and can reach 4 to 10 meters in length; the specimen shown here is about four meters long. (credit a: modification of work by Jan Derk; credit d: modification of work by CDC)

Most free-living flatworms are marine polycladids, although tricladid species live in freshwater or moist terrestrial environments, and there are a number of members from other orders in both environments. The ventral epidermis of free-living flatworms is ciliated, which facilitates their locomotion. Some free-living flatworms are capable of remarkable feats of regeneration in which an individual may regrow its head or tail after being severed, or even several heads if the planaria is cut lengthwise.

The monogeneans are ectoparasites, mostly of fish, with simple life cycles that consist of a free-swimming larva that attaches to a fish, prior to its transformation to the ectoparasitic adult form. The parasite has only one host and that host is usually very specific. The worms may produce enzymes that digest the host tissues, or they may simply graze on surface mucus and skin particles. Most monogeneans are hermaphroditic, but the male gametes develop first and so cross-fertilization is quite common.

The trematodes, or flukes, are internal parasites of mollusks and many other groups, including humans. Trematodes have complex life cycles that involve a primary host in which sexual reproduction occurs, and one or more secondary hosts in which asexual reproduction occurs. The primary host is usually a vertebrate and the secondary host is almost always a mollusk, in which multiple larvae are produced asexually. Trematodes, which attached internally to the host via an oral and medial sucker, are responsible for serious human diseases including schistosomiasis, caused by several species of the blood fluke, Schistosoma spp. Various forms of schistosomiasis infect an estimated 200 million people in the tropics, leading to organ damage, secondary infection by bacteria, and chronic symptoms like fatigue. Infection occurs when the human enters the water and metacercaria larvae, released from the snail host, locate and penetrate the skin. The parasite infects various organs in the body and feeds on red blood cells before reproducing.

Many of the eggs are released in feces and find their way into a waterway, where they are able to reinfect the snail host. The eggs, which have a barb on them, can damage the vascular system of the human host, causing ulceration, abscesses, and bloody diarrhea, wherever they reside, thereby allowing other pathogens to cause secondary infections. In fact, it is the parasite’s eggs that produce most of the main ill effects of schistosomiasis. Many eggs do not make the transit through the veins of the host for elimination, and are swept by blood flow back to the liver and other locations, where they can cause severe inflammation. In the liver, the errant eggs may impede circulation and cause cirrhosis. Control is difficult in impoverished areas in unsanitary, crowded conditions, and prognosis is poor in people with heavy infections of Schistosoma japonicum, without early treatment.

The cestodes, or tapeworms, are also internal parasites, mainly of vertebrates (Figure). Tapeworms, such as those of Taenia spp, live in the intestinal tract of the primary host and remain fixed using a sucker or hooks on the anterior end, or scolex, of the tapeworm body, which is essentially a colony of similar subunits called proglottids. Each proglottid may contain an excretory system with flame cells, along with reproductive structures, both male and female. Because they are so long and flat, tapeworms do not need a digestive system; instead, they absorb nutrients from the food matter surrounding them in the host’s intestine by diffusion.

Proglottids are produced at the scolex and gradually migrate to the end of the tapeworm; at this point, they are “mature” and all structures except fertilized eggs have degenerated. Most reproduction occurs by cross-fertilization between different worms in the same host, but may also occur between proglottids. The mature proglottids detach from the body of the worm and are released into the feces of the organism. The eggs are eaten by an intermediate host, typically another vertebrate. The juvenile worm infects the intermediate host and takes up residence, usually in muscle tissue. When the muscle tissue is consumed by the primary host, the cycle is completed. There are several tapeworm parasites of humans that are transmitted by eating uncooked or poorly cooked pork, beef, or fish.

The tapeworm life cycle begins when eggs or tapeworm segments, called proglottids, pass from human feces into the environment. Taenia saginata infects cattle and Taenia solium infects pigs when they eat contaminated vegetation. The embryo penetrates the animal intestinal wall and takes up residence in muscle tissue, where it transforms into the larval form. Humans who consume raw or undercooked infected meat become infected when the tapeworm attaches itself to the intestinal wall via suckers or hooks on the scolex, or head. The mature worm produces proglottids and eggs, which pass from the body in feces, completing the cycle.
Tapeworm life cycle. Tapeworm (Taenia spp.) infections occur when humans consume raw or undercooked infected meat. (credit: modification of work by CDC)