Nematodes (Roundworms): Classification and Characteristics

What are Nematodes?

Nematodes are also known as roundworms. They are parasites and belong to the phylum Nematoda. They are multicellular animals in the group Ecdysozoa. These nematodes parasitize virtually every type of animal and lots of plants. Roundworms inhabit water bodies and the soil before finding a host. Good topsoil is likely to contain billions of roundworms per acre.

The effect the infestation of these nematodes has on humans, plants, and animals makes the phylum one of the most important of all the parasitic animal groups. Almost every species of invertebrates and vertebrates serve as a host for one or more types of roundworms. The infestation of roundworms in domestic animals causes economical loss. In humans, they cause much discomfort, disease, and at times death.

The predatory species of nematode may feed on small annelids, rotifers, tardigrades, and other nematodes. Many species of nematode penetrate and feed on plant juices from higher plants which cause a great proportion of agricultural damage. The Free-living nematodes feed on yeasts, bacteria, fungi, and algae. They can be saprozoic or coprozoic. Saprozoic meaning that they feed on dead or decaying animal matter. Coprozoic means they can live in fecal material. However, Nematodes themselves can be prey for other organisms like insect larva and mites.

Characteristics of Nematodes

Anatomy

  • Nematodes have a cylindrical shape.
  • They have a bilaterally symmetrical and triploblastic body.
  • Most nematode worms are lesser than 5cm while some parasitic nematodes are more than 1 m in length.
  • Many nematodes are microscopic and some can be seen with your naked eyes.
  • Their outer body is covered by a noncellular cuticle secreted by the underlying epidermis (hypodermis).
Diagram of a nematode as illustrated by Ascaris female.
Diagram of a Nematode as illustrated by Ascaris female showing its structure(A), Cross section(B), and Single muscle cell(C)
Reference: Image from Integrated Principles of Zoology (Fifteenth Edition) Textbook by Hickman, Roberts, Keen, Eisenhour, Larson, I’Anson. Pg 390
  • They shed the cuticle during juvenile growth stages. However, the cuticle is important to nematodes as it helps to contain the high hydrostatic pressure exerted by fluid in the pseudocoel and protects the worm from a hostile environment like the digestive tracts of their host or dry soil.
  • Their hypodermis is syncytial (a large cell-like structure that is formed by joining two or more cells together) and contains dorsal and ventral nerve cords.
  • The body wall muscles of the nematode are longitudinal.
  • They have a hydrostatic skeleton.
  • The body of a nematode is ornamented with ridges, bristles, rings, or other distinctive structures.
A diagram showing the cross section (A) and posterior end (B) of a male Nematode Reference: Image from Integrated Principles of Zoology
A diagram showing the cross-section (A) and posterior end (B) of a Male Nematode
Reference: Image from Integrated Principles of Zoology (Fifteenth Edition) Textbook by Hickman, Roberts, Keen, Eisenhour, Larson, I’Anson. Pg 391

Digestive system

  • The alimentary canal of nematodes consists of a mouth, the pharynx, a short rectum, a long nonmuscular intestine, and a terminal anus.
  • Their pharynx is muscular and has a triradiate lumen.
  • Food is sucked into the pharynx when the muscles in its anterior portion contract rapidly and open the lumen.
  • Relaxation of the muscles anterior to the food mass closes the pharynx lumen. This forces the food posteriorly towards the intestine.
  • The intestine is one-celled thick.
  • The food matter moves posteriorly by body movements and by additional food being passed from the pharynx into the intestine.

Nervous system

  • Around the pharynx, there’s a ring of nerve tissue and ganglia which gives rise to small nerves to the anterior end and two nerve cords, one ventral and one dorsal.
  • There are sensory papillae concentrated around the head and tail of a nematode.
  • They possess amphids which are a pair of more complex sensory organs that open on each side of the head.
  • Amphids are usually reduced in nematode parasites of animals, but most parasitic nematodes bear bilateral pairs of phasmids near the posterior end.

Excretory system

  • Defecation is done by muscles that pull the anus open, and an expelling force is provided by the high pseudocoelomic pressure that surrounds the gut.
  • Their excretory system can be made up of one or more large cells opening by an excretory pore.
  • At times a canal system without glands cells, or both cells and canals together.
  • The excretion of nitrogenous waste is not associated with any specific organ and is passed out in the form of ammonia. 

Reproduction

  • Most nematodes are dioecious (having the male and female reproductive organs in separate individuals).
  • The female nematodes are bigger than the male.
  • Their posterior end bears a pair of copulatory spicules.
  • Fertilization in nematodes is usually internal.
  • Their eggs are stored in the uterus until deposition.
  • When nematodes copulate, one or more Chitinized spicules move out of the cloaca and are inserted into the female genital. The amoeboid sperm then crawl along the spicule into the female.
  • Nematode sperm is the only eukaryotic cell without the globular protein G-actin.

Growth

  • Development among the free-living forms of nematodes is direct. The 4 juvenile stages are separated each by a molt or shedding of the cuticle.
  • Other nematodes require an intermediate host to complete their circle whereas many parasitic nematodes have free-living juvenile stages.

Classification of Nematodes

The classification of nematodes is based on the work of Kampfer, et al. They are classified into two main classes: Adenophorea (Aphasmids) and Secernentea ( Phasmida). Each class is divided into several orders.

ClassOrder
Adenophorea

(Aphasmids)

 

Enoplida

Chromedorida

Dorylaimida

Desmoscolecida

Mermithida

Monohysterida

Araeolaimida

 

Secernentea

(Phasmids)

 

Rhabditida

Strongylida

Spirurida

Oxyurida

Trichuroida

Ascaridida

Camallanida

 

 

  • Class Adenophorea

The nematodes that belong to this class are free-living organisms. The Amphids are generally well-developed. They have about five or more esophageal glands. The excretory system lacks lateral canals. The Caudal glands are present but Phasmid is absent in this class. Some parasites belong to this class. Common examples of nematodes in this class are; Trichinella, Capillaria, Dioctophyme, Trichuris.

However, The class Adenophorea (Aphasmidia) is divided into the following orders:

  • Order Enoplida

They are mostly marine nematodes and are Cyanthiform amphids. Their cuticle contains bristles. A typical example of nematode in this order is Anticoma.

  • Order Chromedorida

This order of nematodes is free-living or marine. Their cuticle lacks any bristles and is smooth or ringed. They have a posterior bulb at the pharynx. Paracanthonchus is a typical example of this order of nematodes.

  • Order Dorylaimida

This order of nematodes is mostly terrestrial. Though it is a diverse order with both freshwater and soil species. The nematode’s buccal cavity consists of a protrusible spear.  It has about 6-10 labial papillae. Their cuticle is without any bristles and smooth. Trichodoris is a typical example of this order.

  • Order Desmoscolecida

This order of nematodes is mostly marine or free-living. Their cuticle is ringed and has prominent bristles with 4 sensory bristles at the anterior end. Desmoscolex is a typical example of Order Desmoscolecida.

  • Order Mermithida

The adult stage of this order is free-living. Although in the larval stage, they live as parasites. The amphids are reduced and their cuticle is smooth. Agamermis and Mermis are typical examples of this order.

  • Order Monohysterida

This order can either be freshwater, marine, or terrestrial nematodes. They have circular amphids. Their cuticle is smooth and ringed with bristles. Monohystera is a typical example of this order.

  • Order Araeolaimida

The amphids of this order are spiral. The nematodes in this order possess labial papillae. Their cuticle may or may not have bristles. However, the cuticle is smooth. A typical example of the order is Plectus.

  • Class Secernentea

The nematodes of this class are mostly parasitic. Though they are both free-living and parasitic. A ventrally coiled phasmid is present in these organisms. They have 3 esophagus glands. However Caudal glands are absent in these nematodes. Their excretory system contains paired lateral canals. Examples of nematodes in the class Secernentea are; Ascaris, Caenorhabditis, Necator, Wuchereria, Enterobius.

The Class Secernentea is divided into the following orders:

  • Order Rhabditida

The nematodes of this order are free-living and parasitic. Their cuticle is ringed and smooth. They have a posterior lobe at the pharynx. The males of this order have copulatory spicules. Rhabditis is a typical example of this order.

  • Order Strongylida

The nematodes of this order are vertebrate parasites. They are devoid of lips and have a well-developed buccal capsule. There is no bulb at their pharynx. They have a true copulatory bursa. A typical example of this order is Strongylus.

  • Order Spirurida

These nematodes are threadlike and vary in size. They can be moderate to large in size. The females can be oviparous or viviparous and are usually larger than males. Their pharynx lacks a posterior bulb. The mouth of these worms has 2 prominent lips. Spirura is a typical example.

  • Order Ascaridida

These nematodes are large and stout. They live as parasites in the intestine of vertebrates. The pharynx of these worms may or may not contain a posterior bulb. They are oviparous. Their mouth contains three prominent lips and they have no buccal capsule. Ascaris is a typical example of this order.

  • Order Oxyurida

These roundworms are usually moderate or small in size and can be vertebrates or invertebrates. Their mouth contains 3-4 simple lips. The males have copulatory spicules. They possess caudal alae. Oxyuris is a typical example.

  • Order Trichuroida

They are commonly called whip-worms. Their pharynx is slender and the mouth lacks lips. Trichuris is a typical example.

  • Order Camallanida

These worms are thread-like and are parasites of aquatic and terrestrial vertebrates. The females are oviparous and the male has no bursa. Camallanus is a typical example.

Nematodes Examples

Examples of Nematodes in Humans and Animals

Ascaris Lumbricoides

Ascaris lumbricoides is one of the most common nematode parasites that is found in humans. In a day, a female Ascaris may lay about 200,000 eggs, which are carried by the host’s feces.

Within 2 weeks under suitable soil conditions, the embryos develop into infective juveniles. Their eggs have a strong tolerance against direct sunlight, high temperature, and desiccation or lack of oxygen.

The shelled juveniles can remain viable for many months or even years in the soil. Ascaris Infection (Ascariasis) happens when the eggs are ingested with uncooked vegetables. Also in some instances when the children put soiled fingers or toys in their mouths.

When the host swallows the embryonated eggs, the eggs eventually hatch and enter into the veins or lymph vessels through burrowing the intestinal wall. When they enter the veins or lymph vessels, they are carried through the heart to the lungs.

There they break out into alveoli, and they are carried up to the trachea.  At this stage, they may cause serious pneumonia if the infection is large. As they reach the pharynx, juveniles are swallowed and pass through the stomach. They mature about 2 months after the eggs were ingested.

The worms cause allergic reactions and abdominal symptoms in the intestine where they feed on intestinal contents. If the worms are in large numbers they can even cause intestinal blockage. Death can occur from Ascaris infection if the intestine is blocked due to heavy infestation.

The infection rate of Ascaris tends to be higher in children and males tend to be more heavily infected than females probably because females practice good hygiene more.

There are other species of ascarid nematodes that are common in wild and domestic animals which is similar to Ascaris. An example is the Toxocara that infects dogs and cats. The life cycle of a Toxocara is similar to an Ascaris. Although the juvenile nematode does not always complete its tissue migration in adult dogs. They remain in the host’s body in a stage of arrested development.

When a female dog is pregnant, it stimulates the juvenile worms to wander and infect the embryo in the uterus. Then when the puppies are born they are born with worms. This is a good reason why pet owners should practice immediate good hygiene disposal of canine wastes.

The Toxocara are also found in humans which leads to larva, a serious condition in children.

Hookworm

These worms are called hookworms because the anterior end curves dorsally, looking like a hook. They have a large plate in their mouth which cuts into the intestinal mucosa of their host. This is where they suck blood and pump it into their intestine, partially digesting it and absorbing the nutrients. They suck a lot of blood and a heavy infection results in anemia in the host. However, hookworm disease in children can cause mental and growth retardation and a general loss of energy.

Lifecycle of Hookworm in Humans
The life cycle of hookworm
The life cycle of Hookworms
Reference: Image from Integrated Principles of Zoology (Fifteenth Edition) Textbook by Hickman, Roberts, Keen, Eisenhour, Larson, I’Anson. Pg 393

The hookworm eggs pass in the feces and the juvenile hatch in the soil. The juveniles feed on bacteria in the soil. The infective juvenile burrow through human skin to the blood when the skin comes in contact with the infected soil. From the blood, it reaches the lungs and finally the intestine.

Trichina worm

Trichinella spiralis is one of several species of tiny nematodes. These nematodes cause Trichinosis, a very lethal disease.

When a poorly cooked or raw meat containing encysted juvenile is swallowed, the worms are liberated into the intestine, where they mature. The adult worms then burrow in the mucosa of the small intestine. This is where the female produces the juvenile.

The juvenile eventually penetrates the blood vessels and is carried throughout the body. They then penetrate skeletal muscle cells. These roundworms are known as one of the largest intracellular parasites.

The Juveniles cause astonishing redirection of gene expression in their host cell, losing its striation and becoming a nurse cell that nourishes the worm.

Trichinella spp. can infect a wide variety of mammals in addition to humans, including hogs, cats, rats, pigs, bears, and dogs. Hogs can get infected by eating garbage that contains pork scraps with juveniles or by eating infected rats.

Pinworm

Pinworms (Enterobius vermicularis) are the most common parasites in the United States. About 30% of children and 16% of adults get infected. Pinworms cause relatively little disease. The adult pinworm lives in the large intestine and cecum.

The females are about 12mm long and move to the anal region to lay their eggs at night. Scratching the resultant itch eventually contaminates bedclothes and hands. The eggs develop fast and become infective within 6 hours of body temperature. Once they are swallowed they hatch in the duodenum and the worms mature in the large intestine.

To diagnose most intestinal roundworms or nematodes, a small bit of feces is examined under the microscope to find their eggs. However, pinworm eggs are usually not found in the feces as the female normally deposits them on the skin around the anus. Hence, the Scotch tape method is more effective. The Scotch tape method involves applying the sticky side of the cellulose tape around the anus to collect the eggs and then placing the tape on a glass slide to examine under a microscope.

Filarial worms

The filarial worm is a parasitic roundworm that causes diseases. At least 8 species of filarial nematodes infect humans. To complete its cycle, this nematode usually requires 2 hosts- an arthropod which is the intermediate host, and a vertebrate which is the primary host. About 120 000 000 people in tropical countries are infected with Wuchereria bancrofti or Brugia malayi.

The filarial worm lives in the lymphatic system and the females are 10cm long. The females release their young ones (tiny microfilariae), into the lymphatic system and blood. As mosquitoes feed, they ingest microfilariae which develops to the infective stage inside the mosquitoes. The microfilariae come out of the mosquitoes when they feed again on humans and penetrates the wound made by the mosquito bite.

After long and repeated exposure to reinfection, a condition known as Elephantiasis manifests.

The condition is seen by excessive growth of connective tissues and enormous swelling of affected parts like the legs, arms, scrotum, and occasionally, the vulva and breast. The disease symptoms are associated with inflammations and obstruction of the lymphatic system.

Another type of filarial worm causes river blindness called Onchocerciasis. The worm is carried by black flies. It infects more than 37 000 000 people in parts of Arabia,  Africa, South America, and Central America.

The most common filarial worm in the united states is likely the Dog heartworm called Dirofilaria immitis.

The heartworm is carried by mosquitoes and infects other animals like cats, sea lions, ferrets, and occasionally humans. Dirofilaria immitis inhabits the hearts, arteries, and lungs of dogs, cats, and some other animals. However, Dirofilaria immitis is more prevalent in dogs. The worm causes a very serious disease among dogs. It is advisable for every dog owner to provide heartworm pills for their dog during mosquito season.

Whipworm

Whipworm is a parasitic nematode called Trichuris trichiura. The nematode is called a whipworm because it is shaped like a whip. One narrow section end of the worm looks like a whip and the other end is thick and resembles a whip handle. Globally, approximately 600- 800 million people have a whipworm infection. This nematode (roundworm) infection can happen in animals too like cats and dogs.

A whipworm infection is known as Trichuriasis. The nematodes infect the large intestine. Once this roundworm reaches the small intestine, its eggs hatch and release its larvae. When the larvae mature, the adult whipworms live in the large intestine. About 2 months later, the female whipworm begins to deposit eggs in the large intestine. They lay about 3000 – 20 000 eggs each day.

Whipworm infection is gotten after consuming dirt or water that has been contaminated with feces containing the whipworm or their eggs. The parasite’s eggs can get into the soil when an infected person/animal defecates outside or probably when the contaminated feces are used as fertilizers. One can unknowingly get infected by touching the dirt and then putting their hands in or near their mouth. At times too by eating fruits or vegetables that have not been thoroughly washed, peeled, or cooked.

The infection often occurs in children. People living in areas with poor sanitation and hygiene and regions with hot, humid climates are susceptible to whipworm infection.

Baylisascaris

Baylisascaris are ascarid nematodes that infect carnivores animals. Some animal groups and species are susceptible to Baylisaascaris larva. Rodents, primates, rabbits, and birds are most often infected. Many cases of infection have been seen in carnivores. However, some species of Baylisascaris are primary to some animals.

For instance, Baylisascaris procyonis infects raccoons, B. columnaris infects skunks, B. melis infects badgers, B. devosi infects martens and fishes. Also B. schroederi infects giant pandas, B. transfuga infects bears, and B. tasmaniensis infects Tasmanian devils, native cats, and quolls.

Baylisaascaris spp. infection occurs when the eggs are ingested directly or the larvae are ingested in a small mammal (intermediate host). When an animal swallows the eggs, the larvae hatch in the intestine, invading the intestinal wall. If in a definitive host, they develop for some weeks and later enter the intestinal lumen where they mature, mate, and produce eggs.

These eggs are then carried out in the fecal stream. For example, an infected raccoon ( the female worm produces about 179 000 eggs in a day) may shed millions of eggs in a day in their feces. The young raccoon may ingest infective eggs and become infected. Whereas older raccoons get infected from third-stage larvae in intermediate (mostly rodents) hosts.

On the other hand,  If the larvae are in a paratenic host they will break into the bloodstream and enter several organs. Most especially, the central nervous system causing great damage. The body tries to respond to the attack by attempting to kill it. In order to escape, the larvae move rapidly in search of the liver, spinal cord, eyes, or brain. At times the larvae can be found in the lungs, heart, and other organs. Later on, the larvae die and the body reabsorbs them.

The animal will not show symptoms of disease If the larvae do not cause any significant damage to important organs of the animal. But if the larvae destroy part of the brain of the host and cause behavioral changes, the host becomes easy prey, bringing the larvae into the new host’s intestine. In small species like mice, it may only take 1 or 2 larvae in the brain of the mice to cause death.

Haemonchus contortus

This nematode is one of the most common pathogens in sheep. It is also known as the barber’s pole worm. The barber’s worm infection is called haemonchosuis. Haemonchus feed on blood and causes anemia, resultant edema, hypoproteinemia, and sometimes death. They are usually found in the abomasums of ruminant animals. Haemonchosis causes huge economic loss for farmers globally. Most especially for farmers in warmer climates.

The animals get infected from ingesting highly infested pastures that contain numerous third-stage larvae. The young sheep ( lamb) are usually susceptible. When the ruminant animals graze, they ingest the Haemonchus larvae on grasses that enter the abomasum. There they may lie inactive within the gastric glands. Once they develop into an adult, they enter into the abomasal surface and attach themselves with a buccal tooth. Then pass eggs in the feces, however,  completing its life cycle.

Anthelmintics are used to treat and prevent this nematode infection and other roundworm infections. Although the resistance of this parasitic nematode against this chemical is growing. However, some ruminant breeds like the  N’Dama cattle and the West African dwarf goat are more resistant to the parasite than other breeds.

Entomopathogenic nematodes

This group of nematodes parasitizes insects.  This roundworm is soft-bodied and non-segmented. They are endoparasitic as they live parasitically inside the infected insect host. Entomopathogenic nematodes are highly diverse, specialized, and complex. They infect many soil insects like the larval form of butterflies, moths, beetles, and flies. Also the adult forms of grasshoppers, beetles, and crickets. They locate their host insects in response to carbon dioxide, other chemical cues, and vibration.

Some entomopathogenic nematodes are used in the biological control of harmful insects and these are the species most studied. Members of Heterorhabditidae and Steinernematidae are such examples. They are the only insect parasitic nematodes that possess an optimal balance of biological control features. The species in these two families (Heterorhabditidae and Steinernematidae) fit perfectly into integrated pest management programs. Most especially because they are seen to be non-toxic to humans.

The juvenile stage of the entomopathogenic nematodes penetrates the host insects through the mouth, spiracles, or anus. In some other species of insects, it penetrates through the intersegmental membranes of the cuticle. After penetrating it enters into the hemocoel. The juvenile stage then releases cells of their symbiotic bacteria from their intestine into the hemocoel of the insects. The bacteria eventually multiply in the hemolymph of the insects causing death within 24-48 hrs. The insect cadaver is usually red if killed by Heterorhabditids and then brown if killed by Steinernematids. The color is due to the pigments produced by the monoculture of the mutualistic bacteria that grows in the host.

Even after the death of the host insects, these nematodes continue to feed on the tissue of the host insects. Hence they mature and reproduce. One or more generations of the nematode may be produced within the host cadaver depending on the resources available. A large number of the infective juvenile produced are then released into the environment to continue their life cycle and infect other insects.

However, reproduction in Steinernematid nematodes and Heterorhabditid nematodes differs. The infective juvenile of Heterorhabditid nematodes eventually become hermaphroditic adults. Although individuals of the next generation produce both females and males. Whereas In Steinernematid nematodes, males and females produce all generations.

Myrmeconema neotropicum

Myrmeconema neotropicum is a tetradonematid nematode parasite known to infect a South American ant (Cephalotes atratus). The South American ant is the only host currently known to be infected by this new species of nematode. The ant has a black abdomen but upon infection, the gaster of the abdomen becomes red, resembling one of the several red berries found in a tropical forest. This nematode induces fruit mimicry.

This nematode’s life cycle begins when a bird eats an infected ant. The nematode eggs are defecated out upon passing through the bird’s digestive system. The defecated eggs are then picked up and fed to the ant larvae( immature ant) by the ant. Once the eggs are inside the larvae’s gut, they migrate to the gaster. It is in the gaster of the ant that they will mature fully.

The mature nematodes begin to reproduce inside the gaster, once the ant larvae pupate. The males of the nematode expire soon after mating whereas the females hold the eggs within themselves. When the ant grows into a young adult, you can easily see the red embryo because the gaster begins to become translucent. The infected ant however travels outside resembling small redberries. Hence being that red berries are bird’s favorite food, the ants are confused for a berry and eaten by birds. Thus starting the whole cycle of Myrrmeconema over again.

Examples of Nematodes in Plants

The nematodes that are parasites of plants are called Eelworms.

Aphelenchoides (Foliar nematodes)

Aphelenchoides are plant-feeding nematodes and are known as Foliar nematodes. They are fungivorus and some species are known to be facultative plant parasites feeding on live plant tissues.

The foliar nematodes infest the aerial portions of the plants, unlike other plant-parasitic nematodes that dwell strictly in soil and roots. The damage they cause to the plants can reduce the yield in food crops and spoil the marketability and appearance of the plants.

Foliar nematodes infest varieties of plants: Clubmosses, ferns, liverworts, angiosperms, gymnosperms, monocots, and dicots.

Vein- delimited discoloration on leaves caused by foliar nematodes
Vein- delimited discoloration on leaves caused by foliar nematodes
Photo credit: www.entnemdept.ufl.edu

Globodera ( Potato cyst nematodes)

Globodera rostochiensis is a plant pathogenic nematode that infects tomatoes and potatoes. This nematode is commonly known as the golden eelworm, yellow potato cyst nematode, or golden nematode.

The nematode infects plants that belong to the family; Solanaceae. They form cysts on the root of the plants. The cysts are yellow in color and are composed of dead nematodes which are usually formed for the female egg’s protection.

When the golden nematode infects the plant, it causes poor growth, wilting, and chlorosis. Further heavy infestation of the nematode can result in water stress, reduced root systems, and deficiency of nutrients. Whereas an indirect infestation may cause an increased vulnerability to fungal infections and premature senescence in the crop.

However, an infestation may take many years to develop and can likely go between 5-7 years unnoticed. The plants may not show unique symptoms during an infestation, hence identifying the parasite is done through soil sample testing. After detecting the nematode, it may take as long as 30 years for the parasite to be effectively and completely eradicated.

Heterodera ( Soybean cyst nematode)

Heterodera glycines is a common nematode known to infest soybean and is known as the soybean cyst nematode. It is a major soybean pest in semi-arid areas of the USA and other places like Brazil, Argentina, Colombia, Egypt, China, Iran,  Japan, Iran,  Korea, Paraguay, and Indonesia.

The eggs of this nematode are usually retained in a cyst formed from the dead female’s cuticle. The infective juvenile in the second stage is found within plant roots and in soil. The mature female is a semi-endoparasite of plant roots and the adult males may be found in the soil.

The yellow dwarf disease of soybean is caused by H. glycines. The infestation causes stunted growth in the plant and can be combined with chlorosis. The infected plants may have yellowed foliage, reduced Rhizobium nodulation, and few lateral roots. The crop yield eventually gets reduced and in worst cases, the plants die.

Meloidogyne (Root-knot nematodes)

Meloidogyne commonly called the root-knot nematode is a pathogenic nematode that attacks the roots of various shrubs, trees, and herbaceous plants. They strive in soil and areas with short winters or hot climates. A lot of plants are susceptible to root-knot nematode infection. These nematodes cause an estimated 5% of global crop loss.

The nematode larvae infect the roots of plants and cause the development of root-knot galls. These galls drain the plant’s nutrients and photosynthate. Young plants infected by the root-knot nematode may die and the infection causes reduced yield in matures plants. Plants that are affected appear discolored, lack vigor, and wilt under stress. There is stunted growth in the plants whose foliage turns yellow and wilts regardless of the adequate soil moisture.

As egg masses, these nematodes survive in soil and in plant debris as infective juveniles. They can be carried by irrigation water to crops. The root-knot nematode at second-stage juvenile infects the plants. This juvenile settles down in the young roots to feed. They later reach maturity after 1-2 months. The female adults are about 0.5-1.0 mm in length and are pear-shaped and translucent. However, they can be found within the galled tissue. Usually, they attach themselves to a gelatinous sac that contains lots of eggs.

Female root-knot nematodes with eggs in galled-roots (microscope view)
Female root-knot nematodes with eggs in galled roots (microscope view).
Photo credit: W. Ye, N.C. Department of Agriculture & Consumer Services. https://content.ces.ncsu.edu

The development of the parasite in some cases is parthenogenic. In others, before eggs are laid, the tiny worm-like male mates with the female. The first stage juvenile then develops within the eggs. Then, the juveniles break free at the second stage and either migrate inside the root or escape into the soil to commence feeding some other place. It’s been observed that these nematodes in dry conditions survive not more than a few weeks but can survive for about 3 months in moist soil.

Nacobbus

Nacobbus is a plant-parasitic nematode that attacks crops like tomatoes, sugarbeet, potatoes, and quinoa. There are 3 species of the genus Nacobbus: N. aberrans, N. dorsalis, N. bolivianus. These nematodes are prevalent in South and North America. N. abberans is a common pest of potatoes in South America and of sugarbeet in North America while N. dorsalis is somewhat rare and seen mainly in some locations in California.

The juvenile Naccobus is a migratory endoparasite that penetrates the roots tips or axial roots of the plants. This penetration causes slight swelling on the roots at their axis and tip. Hence inducing the formation of enlarged cells and root galls.

However, the swellings on sugarbeets may extend over a huge portion of the root axis, and the potato roots after being invaded by the juvenile may exhibit lesions with discolored tissues.

Pratylenchus (Lesion nematodes)

Pratylenchus is commonly known as lesion nematodes. They are plant-parasitic and are known for the root-lesion disease in most crops like soybeans, corn, banana, wheat, and potatoes. However, Pratylenchus is mostly found in corn. These nematodes are migratory endoparasites. They move around, feed, and reproduce in the root of the host plants, feeding mainly on the cortex of the host root.

In many species of Pratylenchus, males are rare. However, the female root-lesion nematode lays single eggs in the root of the plant or in the soil. They usually reproduce through parthenogenesis, which involves laying eggs without fertilization.

Necrotic lesions of the host plant roots are a major symptom of the disease caused by the nematode. A secondary infection can also occur as the lesion may serve as an entrance for pathogenic fungi and bacteria to cause another infection.

Lesion nematode damage on roots
Roots damaged by lesion nematode will become dark reddish-brown to black, and necrotic (rotting).
Here, healthy roots and heavily infected roots can be seen in a cover crop.
Healthy roots are usually pale white to tan.
Photo Credit: A. Gorny, NC State University
https://content.ces.ncsu.edu

The infected plant becomes stunted in growth. Also, the crop becomes chlorotic and wilted. In most cases, the crop dies. Hence the root-lesion nematode causes a significant yield loss.

Trichodorus obtusus

Trichodorus obtusus is a plant-parasitic nematode commonly called stubby root nematodes. They are named so because they feed and cause a stunted and stubby appearing root system in the host plant. The stubby-root nematode is an ectoparasitic nematode that feeds on the meristem cells of the plant root tip as their bodies remain in the soil.

These nematodes attack turf grasses and other plants like bermudagrass, zoysiagrass, tomato, Sideoats grama, eucalyptus, Saw palmetto, potato, Kentucky bluegrass, rhododendron, big bluestem, seashore paspalum, sorghum-sudangrass, sweetbay magnolia, little leaf linden, and St. Augustine grass.

Stubby root nematodes are characterized by possessing a six-layer cuticle and are unique among other nematodes for having an onchiostyle, a curved solid stylet, or spear which they use for feeding. They make use of the onchiostyle like a dagger to puncture holes in plant cells whereas other plant-parasitic nematodes possess straight, hollow stylets.

After puncturing the plant cells, the nematode then secrets salivary material from its mouth into the punctured plant cell. This salivary material hardens and forms a feeding tube. The nematode then uses the feeding tube as a straw to withdraw and ingest the cell contents of the host plant. After feeding on an individual cell, the nematode moves on to other cells to feed.

They leave the old feeding tube behind and form new ones in each cell that they feed on. The stubby root nematode eventually causes stunted roots and also transmits viruses. The Common viruses these nematodes transmit are tobravirus, pea early-browning virus, tobacco rattle virus, and pepper ringspot virus.

Trichodorus obtusus have to mate in order to produce offsprings. However, the female Stubby root nematodes lay eggs after mating. The eggs remain in the soil until they hatch. The eggs hatch as second-stage juveniles. As they must feed on plants to reproduce and survive. When they locate a root and commence feeding, the juvenile nematode will molt 3 times before becoming an egg-laying adult nematode.

The nematode damages the root system of its host plants making it prone to environmental stresses. This can lead to an increase in the use of water and fertilizer inputs. It also makes the plants less competitive with weeds leading to an increase in the use of herbicides.

For instance, the damage caused by Trichodorus obtusus on turf grasses usually occurs in irregularly shaped patches within a given area. The turfgrass may wilt in those areas, thin out, and eventually, die if drought occurs. The symptoms however are more severe in sandy soil than heavier soils.

Xiphinema  (Dagger nematode)

These nematodes parasitize plants and are also known as dagger nematodes. They are ectoparasites and all stages (except eggs) of these nematodes attack and feed on the host plant’s root. Major species of this nematodes include X.italiae, X.americanum, X.diversicaudatum, X.pachtaicum, and X.index. They are characterized by their long bodies and stylet.

The stylet is usually long enough to reach the plant’s vascular tissue. As the body of the dagger nematode stays in the soil outside the plant root, it inserts its long stylet deep into the root. The stylet penetrates the plant tissues and punctures cell walls. As they feed, they secrete enzymes to digest the plant cell content. These enzymes digest and destroy the root cells which cause a malformation of the root tissues. Eventually, the root cells collapse.

Different members of this nematode group through penetration have been known to cause a moderately large amount of damage to the plant roots, resulting in gall formation in some species. Xiphinema attacks a large variety of plants like grape, strawberry, hops, nectarine, oak, raspberry, grapevine, carrot, peach, cherry, rose, and soybean.

Species of Xiphinema are sensitive to soil moisture and temperature changes. Hence they migrate away from desiccating conditions in topsoil. Therefore, most of them can survive and live deep in the soil.

Dagger nematodes are vectors of nepoviruses. Most of these nematodes during feeding transfer viruses to plants. Some of the viruses they transfer are the Tomato ringspot virus, cherry rasp leaf virus, and tobacco ringspot virus. They feed on the host plant’s roots and spread viral mosaic and wilting diseases, which can cause economic damage and death. Controlling viral disease in susceptible crops is a major challenge. However, studies have shown that biofumigation and crop rotation are effective Control measures to an extent.

Xiphinema has 6 life cycle stages. The females lay their eggs singly in thin water layers in the soil. There are 3 or 4 molts after the first stage juvenile emerges from the egg. With each molt, they increase in size until they become adults. While the vector-capable juveniles feed on the virus-infected plants, they mature into adults. Also, they acquire plant pathogenic viruses that form a lining in the pharynx-stylet tube. During feeding, the viruses are then injected into the root tissues. The feeding at the meristematic root tip causes damage. It destroys the root cells and causes a reduction in the root volume. Also, causing an above-ground effect of stunted growth of the crop and patchy fields.

Bursaphelenchus

Bursaphelenchus is a genus of nematodes that cause severe wilt diseases in a number of tree species. There are two species of this roundworm; the pinewood nematode (B. xylophilus) and the red ring nematode (B. cocophilus). However, these nematodes have built a symbiotic relationship with some insects such as weevils and beetles.

They invade these insects and are transported from a diseased tree to a healthy tree. There are two common tree wilt diseases caused by these nematodes (i.e the pine wilt and coconut red ring disease). They are significant pests of pine trees and coconut palms.

The pine wilt disease is caused by B. xylophilus and beetles of the genus Monochamus are vectors of the disease.

B. cocophilus causes the coconut red ring disease and has two vectors- the sugarcane weevil Metamasius sp and the palm weevil Rhynchophorus sp

These nematodes inhabit decaying wood or soil. They feed on fungal hyphae growing on the wood or feed on the wood itself. When they reduce the fungal load on the wood, they are said to be beneficial.  However, as they consume the tissue of the plant, they can make the tree die.

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