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DESCRIPTION OF THE PHYLUM NEMATODA

DESCRIPTION OF THE PHYLUM NEMATODA (DIESING 1861)

EUKARYA> UNIKONTA> OPISTHOKONTA> ANIMALIA> BILATERIA> PROTOSTOMATA> ECDYSOZOA> NEMATOIDA> NEMATODA
Nematoda (ne-ma-TO-da) is formed from the Greek word for thread -nema (νήμα). The reference is to the long, thread-like appearance of the animals, especially because they have no obvious external structures other than a smooth cuticle. Alternative name for the phylum is Nemata.
INTRODUCTION TO THE NEMATODA

The nematodes are among the most ubiquitous of animals, and they are also about the most difficult to differentiate. Figure 1 shows the general vermiform appearance of all members of the group. Thus, without doubt, there are many more than the 15,000+ described species. Likely, the number of species could be more than a million (Nielsen 2001). Most are free-living, but many are some of the most important (economically important) parasites of our species and of our domesticated plants and animals.

Nematodes have only a few obvious characters other than the extreme reduction to a worm-like animal. Amphids, sensory structures located around the anterior end, and a unique excretory system appear to be true synapomorphies (Pechenik 2005). Otherwise, they have an odd set of characters, especially when comparing them to their likely cohort which includes the Loricifera, Kinorhyncha, Priapulida, and Nematomorpha. They seem to have lost the segmentation of most of the taxa as well as the circular muscles, and the introvert.

Nematodes have only longitudinal muscle lining the pseudocoelom, which gives them a characteristic stiff undulating motion (Figure 1). The muscle also sends extensions to the nerve cords (rather than the norm of having axons travel to the muscle). Also, the sexes are separate and dimorphic. The females are always larger. The males usually have a a slight curl to the end of tail and often have papillae and other lateral extensions on the posterior end. The mouth is terminal and surrounded by three lips (Figure 2). The anus is subterminal.

Typically, nematodes have a fairly simple life cycle that involves at least four molts from the egg to the adult. The four larvae (referred to as L1 to L4) have fairly distinctive and universal characteristics. The first two larval forms usually have a muscular pharynx. At the second molt, the outer cuticle remains around the third stage larva (L3), particularly in parasitic forms.

Giribet et al. (2007) confirm that the nemtodes are a natural group and a sister group to the Nematomorpha in a clade called the Nematoida (a term coined by Schmidt-Rhaesa 1998) which includes Nematoda and Nematomorpha, phyla that share obvious morphological similarities. Furthermore, Telford et al. (2008) list 5 morphological synapomorphies, first identified by Nielsen (2001). The molecular support, though present, is weak (Telford et al. 2008, Petersen and Ernesse 2001, and Mallatt et al. 2003).

They fall within a clade of protostome animals called the Ecdysozoa (Nicholas 2001b), a group that Nielsen (2001) does not recognize as valid. In fact, Nielsen (2001) suggests that the nematodes could be highly reduced insects. Nevertheless, much strong evidence for the monophyly of the Ecdysozoa has emerged (Telford et al. 2008, Philippe et al. 2005, and Webster et al. 2006).

The system of Brusca and Brusca (2003) divides the nematodes into two large groups (Figure 3) according to the occurrence of phasmids, sensory organs near the posterior ends of many parasitic nematodes. Adenophora (also called Aphasmida) do not have phasmids and are mostly free-living. Secerentea (also called Phasmida) have phasmids, and many of the nematode parasites of human concern are from this group. Blaxter et al. (1998) and Dorris et al. (1999) convincingly demonstrate that although the Secerentea (Phasmida) is monophyletic, the Adenophorea (Aphasmida) is paraphyletic and should be separated into at least three taxa of equal rank. So, until the systematics of nematodes settles out, this will be a tentative classification.
FIGURE 1. A nematode viewed with the SEM.
Image from http://www.ucmp.berkeley.edu/phyla/ecdysozoa/nematode.jpg
FIGURE 2. The anterior end of an Ascaris showing the three “lips” over the mouth. Image taken with an SEM.
Image from http://www.biosci.ohio-state.edu/~parasite/ascaris.html

FIGURE 3. The two classes of Nematoda (in shaded box) in the context of the Nematoida (N) and other phyla of the Ecdysozoa (clade E).

P = Protostomata

E = Ecdysozoa

S = Scalidophora

N = Nematoida

PA = Panarthropoda

ADENOPHOREA

The Adenophorea (also called the Aphasmida) seems to be the most primitive group of nematodes. Mainly, they are free-living in soil and water; however, there are a few parasitic forms of aphasmids. As the alternate name implies, they do not have phasmids (text with tooltip) A phasmid is a sensory organ of parasitic nematodes. , and the amphids (text with tooltip) Amphids (n.) are chemosensory organs at the anterior end of some Nematoda. are located posteriorly on the head region. In fact, they have no sensory bristles or papillae on the head and body. They are simple, spindle-shaped worms with simple excretory organs (single-celled).

  • Several taxa of economic importance include Trichinella (Trichina Worm, Figure 4; see the Life Cycle of Trichinella) and Trichuris (Whipworm, Figure 5; see the Life Cycle of Trichuris). Both of them have relatively simple life cycles. Trichinella lives as small adults in the gut. Females brood the eggs and L1 larvae in their bodies and release them after one molt as L2 larvae that burrow into the intestinal mucosa, get into the circulatory system and travel to the tissue where they encapsulate as an L3 larva. Mostly, they inhabit striated muscle (Figure 3), especially the diaphragm. However, they can be found in almost all tissue, including the brain. They may even travel across the placenta and infect a developing fetus. Usually, in humans the cycle ends there, but in animals that practice cannibalism or prey on mammals that might be infect, the L3 larva travels through the stomach and emerges from its cyst in the small intestine. There, they molt and copulate.
FIGURE 4. Trichinella encapsulated as an L3 larva in muscle tissue.
Image from http://elegans.swmed.edu/Nematodes/
FIGURE 5. Female (L) and male (r) Trichuris. Note the difference in size and the long whip-like extention of the animal.
Image from http://www.stanford.edu/…/Trichuris/Untitled-12.htm

SECERENTEA

Secerentea (Phasmida) is a monophyletic group defined by the presence of phasmids. Amphids are located anteriorly in the head region. The excretory system is more complex and includes collecting ducts. Males very often have setae or papillae on the tail, where they also might have lateral extensions. Members of this group include some of the most destructive parasites and include Ascaris, Necator (Hookworm), Dracunculus (Guinea Worm), and Wuchereria (Microfilaria).

  • Ascaris is an intestinal parasite that can grow to 50 cm. The female continually releases eggs that pass out of the digestive tract with the feces. The eggs can survive extreme conditions, including those of a sewage treatment plant. Indeed, when I was a graduate student, someone in our group was diagnosed with Ascariasis, which she likely contracted by handling preserved animals. Furthermore, when human feces are used as fertilizer, the incidence of Ascariasis in the community can approach 100%. The eggs have a mucilaginous outer covering which can stick to developing plants. If the food being grown is eaten raw, the eggs go through the stomach, hatch, and the L1 larva emerges. It burrows through the gut wall, gets into the circulatory system and travels to the lungs where it molts and grows in the alveolar space. It molts again and induces the host to cough. The larva is thus brought up to the throat and swallowed. Back in the small intestine, the larva molts once more and grows. Infestations can be very high (Figure 6). Noble and Noble (1976) report that one man had as many as 1,488 worms when he was autopsied. See the Life Cycle of Ascaris
  • Necator, the Hookworm, as a mature adult has three razor-like lips with which it slices the intestinal mucosa and pulls the bleeding portion into its buccal area (Figure 7). In that way it feeds on the blood as it oozes from the injured mucosa. Females release eggs which are carried out of the body with the fecal material and hatch on the ground. The larvae are free-living and molt twice to become infective L3 larvae, which burrow into the skin of a host and then travel to the lungs to complete the lifecycle in the same way as Ascaris. See the Life Cycle of Necator
  • Dracunculus (the Guinea Worm) lives just under the skin and can grow to a length of more than a meter. The female broods the young in her body and when a host steps into water, she sticks her posterior end out through a hole that she maintains as an open sore. She can expel hundreds of larvae which infect microcrustacea where they live in the haemocel and molt to become infective. The primary host acquires a Guinea Worm by drinking water and ingesting an infected microcrustacean. Larvae burrow through the intestinal wall and develop in the connective tissue where copulation occurs. Males die and the females migrate to the subcutaneous tissue of a lower extremity to complete the cycle. The worms are removed by putting the leg into water and catching the worm as it emerges to release larvae. Once caught, the worm is attached to something like a matchstick and slowly rolled up until it is removed (see Figure 8). See the Life Cycle of Dracunculus
  • Filarial worms (Figure 9), like Brugia, live their lives in the lymphatic system. Larvae are released and travel into the peripheral blood, where they may be taken by a mosquito vector. They mature in the mosquito and infective larvae then are injected into another host when she takes another blood meal. Filiariae can pack lymph nodes and cause fluid to pool in the extremities to produce a condition called elephantiasis. See the Life Cycle of Brugia

As debilitating as some of these worms might be, their greatest impact is on the loss of agricultural production (livestock and plants) which have been infected. Whole crops like tomatoes can succumb completely to a heavy infestation in which infective larvae penetrate the roots and cause galls to develop which leads to stunting, low yield, and early die off. Similar problems occur in corn, wheat and rice, all major sources of calories for humanity. Even large trees, like pecans, are plagued by certain nematode parasites.

FIGURE 6. A group of Ascaris passed as a bolus from an infected human.
Image from http://www.biosci.ohio-state.edu/~parasite/ascaris.html
FIGURE 7. SEM micrograph of the head of Necator. Note the razor-like teeth that are used to slice into the intestinal mucosa.
Image from http://instruction.cvhs.okstate.edu/JCF … g0070a.jpg
FIGURE 8. The careful removal of a Guinea Worm, Dracunculus from the foot of a person who is infected.
Image from http://ib.berkeley.edu/courses/ib116/guinea-worm.jpg
FIGURE 9. A filarial worm (Brugia) in a blood smear. These can become so abundant that they block the lymph nodes and cause a diseace called Elephantiasis.
Image from the CDC and in the Public Domain

Most nematodes likely are free-living and frequently seen in water and soil samples in high densities. Certain fungi take advantage of this source of food and produce loop traps in their mycelia (text with tooltip) A mycelium (mycelia, pl.) is a mass of hyphae or fungus-like filaments. . Upon sticking its head through the loop, a nematode will find itself caught by the sudden swelling of the fungal filaments (Figure 10). The fungus then feeds on the dying worm.

FIGURE 10. An SEM micrograph of a free-living soil nematode being caught by a soil fungus.
Image from http://www.extension.umn.edu/distribution/cropsystems/images/M1272-7.jpg
LITERATURE CITED

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By Jack R. Holt and Carlos A. Iudica. Last revised: 02/03/2014
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