Describe the structure and systems of flatwormsDescribe the structural organization of nematodesCompare the internal systems and the appendage specialization of arthropods

The animal phyla of this and subsequent modules are triploblastic and have an embryonic mesoderm sandwiched between the ectoderm and endoderm. These phyla are also bilaterally symmetrical, meaning that a longitudinal section will divide them into right and left sides that are mirror images of each other. Associated with bilateralism is the beginning of cephalization, the evolution of a concentration of nervous tissues and sensory organs in the head of the organism, which is where the organism first encounters its environment.

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The flatworms are acoelomate organisms that include free-living and parasitic forms. The nematodes, or roundworms, possess a pseudocoelom and consist of both free-living and parasitic forms. Finally, the arthropods, one of the most successful taxonomic groups on the planet, are coelomate organisms with a hard exoskeleton and jointed appendages. The nematodes and the arthropods belong to a clade with a common ancestor, called Ecdysozoa. The name comes from the word ecdysis, which refers to the periodic shedding, or molting, of the exoskeleton. The ecdysozoan phyla have a hard cuticle covering their bodies that must be periodically shed and replaced for them to increase in size.

Flatworms

The relationships among flatworms, or phylum Platyhelminthes, is being revised and the description here will follow the traditional groupings. Most flatworms are parasitic, including important parasites of humans. Flatworms have three embryonic germ layers that give rise to surfaces covering tissues, internal tissues, and the lining of the digestive system. The epidermal tissue is a single layer of cells or a layer of fused cells covering a layer of circular muscle above a layer of longitudinal muscle. The mesodermal tissues include support cells and secretory cells that secrete mucus and other materials to the surface. The flatworms are acoelomate, so their bodies contain no cavities or spaces between the outer surface and the inner digestive tract.

Physiological Processes of Flatworms

Free-living species of flatworms are predators or scavengers, whereas parasitic forms feed from the tissues of their hosts. Most flatworms have an incomplete digestive system with an opening, the “mouth,” that is also used to expel digestive system wastes. Some species also have an anal opening. The gut may be a simple sac or highly branched. Digestion is extracellular, with enzymes secreted into the space by cells lining the tract, and digested materials taken into the same cells by phagocytosis. One group, the cestodes, does not have a digestive system, because their parasitic lifestyle and the environment in which they live (suspended within the digestive cavity of their host) allows them to absorb nutrients directly across their body wall. Flatworms have an excretory system with a network of tubules throughout the body that open to the environment and nearby flame cells, whose cilia beat to direct waste fluids concentrated in the tubules out of the body. The system is responsible for regulation of dissolved salts and excretion of nitrogenous wastes. The nervous system consists of a pair of nerve cords running the length of the body with connections between them and a large ganglion or concentration of nerve cells at the anterior end of the worm; here, there may also be a concentration of photosensory and chemosensory cells (

).

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Figure 2: Phylum Platyhelminthes is divided into four classes: (a) Bedford’s Flatworm (Pseudobiceros bedfordi) and the (b) planarian belong to class Turbellaria; (c) the Trematoda class includes about 20,000 species, most of which are parasitic; (d) class Cestoda includes tapeworms such as this Taenia saginata; and the parasitic class Monogenea (not shown). (credit a: modification of work by Jan Derk; credit c: modification of work by “Sahaquiel9102″/Wikimedia Commons; credit d: modification of work by CDC)

The monogeneans are external parasites mostly of fish with life cycles consisting of a free-swimming larva that attaches to a fish to begin transformation to the parasitic adult form. They have only one host during their life, typically of just one species. The worms may produce enzymes that digest the host tissues or graze on surface mucus and skin particles. Most monogeneans are hermaphroditic, but the sperm develop first, and it is typical for them to mate between individuals and not to self-fertilize.

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 almost always a mollusk. Trematodes are responsible for serious human diseases including schistosomiasis, caused by a blood fluke (Schistosoma). The disease infects an estimated 200 million people in the tropics and leads to organ damage and chronic symptoms including fatigue. Infection occurs when a human enters the water, and a larva, released from the primary snail host, locates and penetrates 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 primary snail host.

The cestodes, or tapeworms, are also internal parasites, mainly of vertebrates. Tapeworms live in the intestinal tract of the primary host and remain fixed using a sucker on the anterior end, or scolex, of the tapeworm body. The remaining body of the tapeworm is made up of a long series of units called proglottids, each of which may contain an excretory system with flame cells, but will contain reproductive structures, both male and female. Tapeworms do not have a digestive system, they absorb nutrients from the food matter passing them in the host’s intestine. Proglottids are produced at the scolex and are pushed to the end of the tapeworm as new proglottids form, at which point, they are “mature” and all structures except fertilized eggs have degenerated. Most reproduction occurs by cross-fertilization. The proglottid detaches and is released in the feces of the host. The fertilized eggs are eaten by an intermediate host. The juvenile worms emerge and infect the intermediate host, taking up residence, usually in muscle tissue. When the muscle tissue is eaten by the primary host, the cycle is completed. There are several tapeworm parasites of humans that are acquired by eating uncooked or poorly cooked pork, beef, and fish.

Nematodes

The phylum Nematoda, or roundworms, includes more than 28,000 species with an estimated 16,000 parasitic species. The name Nematoda is derived from the Greek word “nemos,” which means “thread.” Nematodes are present in all habitats and are extremely common, although they are usually not visible (

).

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Figure 3: (a) An scanning electron micrograph of the nematode Heterodera glycines and (b) a schematic representation of the anatomy of a nematode are shown. (credit a: modification of work by USDA, ARS; scale-bar data from Matt Russell)

Most nematodes look similar to each other: slender tubes, tapered at each end (

). Nematodes are pseudocoelomates and have a complete digestive system with a distinct mouth and anus.

The nematode body is encased in a cuticle, a flexible but tough exoskeleton, or external skeleton, which offers protection and support. The cuticle contains a carbohydrate-protein polymer called chitin. The cuticle also lines the pharynx and rectum. Although the exoskeleton provides protection, it restricts growth, and therefore must be continually shed and replaced as the animal increases in size.

A nematode’s mouth opens at the anterior end with three or six lips and, in some species, teeth in the form of cuticular extensions. There may also be a sharp stylet that can protrude from the mouth to stab prey or pierce plant or animal cells. The mouth leads to a muscular pharynx and intestine, leading to the rectum and anal opening at the posterior end.

Physiological Processes of Nematodes

In nematodes, the excretory system is not specialized. Nitrogenous wastes are removed by diffusion. In marine nematodes, regulation of water and salt is achieved by specialized glands that remove unwanted ions while maintaining internal body fluid concentrations.

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Most nematodes have four nerve cords that run along the length of the body on the top, bottom, and sides. The nerve cords fuse in a ring around the pharynx, to form a head ganglion or “brain” of the worm, as well as at the posterior end to form the tail ganglion. Beneath the epidermis lies a layer of longitudinal muscles that permits only side-to-side, wave-like undulation of the body.