HISTORY AND DEVELOPMENT OF NEMATODE

World history and development:

• Turbevill Needham (1743) - He crushed shrunken, blackened wheat grain and
examined nematodes under primitive microscope. Then it was named as Vibrio tritici
(=Anguina tritici) by Steinbuch in 1799. It is First plant parasitic nematode report in
International level.
• Berkeley (1855)- Observed Vibrios from galls on the root of cucumber plants in
greenhouse in England and it was named as of root knot nematode, Meloidogyne spp.
by Cornu 1879.
• Kuhn (1857) - Reported the stem and bulb nematode, Ditylenchus dipsaci infesting the
heads of teasel.
• Schacht (1859) - Report of sugarbeet cyst nematode, Heterodera schachtii (named by
Schmidt in 1871) from Germany.
• Atkinson & Neal (1889)- Publication about the root knot nematodes in the United
States.
• N.A. Cobb (1907) - Joined in the United States Department of Agriculture (USDA) and
considered to be the Father of American Nematology
• N.A. Cobb (1914-1918)- Contributions to the Science of Nematology, Development of
methods and apparatus used in Nematology.
Contribution of N.A.Cobb
✓ 1907- He joined the USDA and considered to be the Father of American
Nematology.
✓ 1913-Published ‘Contribution to Science of Nematology.
Some important contribution of N.A.Cobb
✓ Developed several techniques which are routinely used today, like Sieving method
for nematode extraction from soil, mounting of nematodes on double coverslip
alluminium slide, preparation of en-face views, new camera lucida apparatus for
making nematode illustrations etc
✓ Description of several minute sensory organs like amphids, deirids, cephalids,
papillae etc in nematodes
✓ Established separate Division of Nematology at USDA
✓ Separation of free living and plant parasitic nematodes from Helminthology to
nematology (term proposed during 1914).
• Filipjev (1941)- Discovery of potato cyst nematode Globodera rostochiensis from
USA.
• 1955- European Society of Nematologists founded
• Canada (1955)- Peach replant problems by lesion nematode, Pratylenchus penetrans.
Mountain and Patrick first time demonstrated the pathogenicity of plant parasitic
nematodes to prove Koch’s postulates.
• Hewitt, Raski and Goheen (1958) - First report of transmission of viruses by
nematodes (Xiphinema index transmitting ‘fan leaf virus’ in grapevine in USA.
• 1961- Society of Nematologists founded in the United States.
 • 1969 - The Nematological Society of India (NSI) was established.
Indian history and development:
• C.A.Barber (1901) - Root knot nematode on tea in Devala estate, Tamil Nadu, South
India. First ever report of a plant parasitic nematode from India.
• Butler (1906) - Reported root knot nematode on black pepper in Kerala.
• Butler (1913-1916)- Reported Ufra disease of rice due to the infestation of Ditylenchus
angustus in Bengal.
• Ayyar (1926-1933) - Reported root knot nematode infestation on vegetable and other
crops in South India.
• Dastur (1934-1936)– Reported white tip disease of rice caused by Aphelenchoides
besseyi in Central India.
• M.R.Siddiqui (1959-1961)- Recorded citrus nematode – Tylenchulus semipenetrans
from Uttra Pradesh.
• 1961 – Nematology laboratory established at AC &RI, Coimbatore under Rock feller
foundation and ICAR by Dr. A.R. Seshadri. Jones found Golden nematode of potato–
Globodera rostochiensis first time in the Nilgiris hill of Tamil Nadu Nematology unit
established at IARI, New Delhi – Nematology unit established at the Central Potato
Research Institute, Shimla.
• 1963 – Laboratory for potato cyst nematode research established at Uthagamandalam
with the assistance of Indian Council of Agriculture Research.
• 1964 – First International Nematology course held at AMU, Aligarph and IARI, New
Delhi in collaboration with International Agriculture College, Netherland.
• Nair, Dass and Menon (1966)- reported the burrowing nematode, Radopholus similis
on banana for the first time from Kerala.
• 1969- Nematological Society of India founded and first All India Nematology
Symposium held at IARI, New Delhi.
• 1971 - Indian Journal of Nematology was published.
• 1971 - Fourth South East Asian Nematology course at New Delhi.
• 1977 - Started All India Co-ordinated Research Project (AICRP) on Nematode pests of
crops and their control sponsored by DST. Subsequently taken by ICAR, New Delhi in
1979. Now functioning in 18 centres in all over India with its Project Co-ordinator at
IARI, New Delhi.
• 1979- M.Sc. (Ag.) Plant Nematology course started at Tamil Nadu Agricultural
University, Coimbatore.
• 2000 - National Nematology Symposium on “Integrated Nematode Management” held
at OUAT, Bhubaneshwar, Odisha. 2001 National Congress on “Centenary of
Nematology in India: Appraisal and Future plans” at IARI, New Delhi.
• 2015 - Department of Nematology established at Kerala Agriculture University,
Trivandrum
 DEFINITION AND ECONOMIC IMPORTANCE OF NEMATOLOGY
Nematology - The study of plant parasitic nematodes and free-living nematodes is called
Nematology.
Nematode - Nematodes are triploblastic (having three germ layers constituting the embryo),
bilaterally symmetrical, unsegmented, Pseudocoelomate (body cavity without epithelium),
colorless and verimiform animals.The word ‘Nematode’ is derived from Greek words - ‘nema’
means thread, and ‘oides’ or ‘eidos’ means resembling of form, thus nematodes are thread like
forms.
Economic importance of nematodes:
India’s top ten important plant parasitic nematode genera are Meloidogyne, Heterodera,
Anguina, Radopholus, Pratylenchus, Tylenchulus, Rotylenchulus, Hirschmanniella,
Tylenhcorhynchus and Ditylenchus
1. Seed gall nematode, Anguina tritici responsible for ‘ear cockle’ diseases of wheat in
Northern India cause yield loss about 10,000 tonnes with 70 million rupees.
2. Cereal cyst nematode, Heterodera avenae causes the ‘molya’ disease of wheat and barley
in Rajasthan, Punjab, Haryana, Himachal Pradesh and Jammu and Kashmir. The loss due to
this nematode is about 32 million rupees in wheat and 25 million rupees for barley in Rajasthan
State alone. For Haryana it causes 688 million rupees (based on MSP).
3. Golden cyst nematode of potato Globodera rostochiensis is a serious problem in the hills
of Nilgiri and Kodaikanal cause 12-25% loss. Under server infestation total failure of crop also
recorded.
4. An annual loss of 20 million was assessed in coffee due to lesion nematode, Pratylenchus
coffeae in an area of 1000 ha in Karnataka state alone. It causes foot rot of young plants and
decline or die back of older plants.
5. The burrowing nematodes, Radopholus similis cause yield loss in black pepper, arecanut
and banana. It causes slow wilt of pepper and toppling disease in banana.
6. The citrus nematode, Tylenchulus semipenetrans causes slow decline in citrus.
7. The reniform nematode, Rotylenchulus reniformis attacks a large number of plants and
cause considerable loss to vegetables and pulses.
8. The root knot nematode, Meloidogyne species produced spectacular symptom of galls
produced on roots of vegetables, fruits, flower, and ornamental crops.
 GENERAL CHARACTERISTICS OF PLANT PARASITIC NEMATODE
1. The nematodes possess elongate, unsegmented, cylindrical or worm like body tapering
towards both ends, unciliated and circular in cross section.
2. Body is bilaterally symmetrical.
3. They are aquatic, terrestrial and parasitic or free living.
4. The body is covered by tough and resistant cuticle secreted by epidermal (hypodermal)
cells.
5. Terminal oral aperture (mouth) surrounded with lips and papillae.
6. Digestive system consists of feeding apparatus, oesophagus, intestine and rectum.
7. Body consists of two tubes.
8. The nervous system consists of circum-oesophageal nerve ring and longitudinal nerves.
9. Primitive excretory system is devoid of protonephridial cilia (is an excretory tube that
lacks an internal opening) or matanephridial funnel (is an excretory organ found in
invertebrates like arthropods, annelids and molluscs).
10. The circulatory and respiratory systems completely absent.
11. The females have separate genital pore and males have a common opening known
cloaca and well-developed copulatory apparatus consisting of spicules and
gubernaculum.
12. Females are oviparous or ovoviviparous or viviparous (that produce live babies from
their bodies rather than eggs). The cleavage is terminated and growth is accompanied
by molting.
13. Life cycle is direct and there are four juvenile stages.
 MORPHOLOGY OF NEMATODES
The size of nematode may vary from 0.2 mm to about 12 mm length with an average
of 0.01 mm and 0.5 mm in breadth (1 to 15 % to length). Males are smaller than females.The
body of nematode is tubular which may be divided into three regions
1. The outer body tube or body wall
2. Inner body tube – Digestive system
3. Body cavity or pseudocoelome – Reproductive system, Nervous system, Excretory
system
1. Outer body tube:
The outer body tube or body wall includes the cuticle or exoskeleton,
hypodermis and somatic muscles. The outer body tube or body wall includes the cuticle,
hypodermis and somatic muscles. The body wall protects the nematode from the harsh
external environment, serves as the exoskeleton and provides the mechanism for
movement of the organism through the soil and plant tissue.
a) Exoskeleton or cuticle- The cuticle is a non-living, non-cellular, triple-layered
covering that is secreted by the underlying hypodermis. The cuticle is flexible. It
covers the entire body and lines the oesophagus, vulva, anus, cloaca, excretory pore
and sensory organs. The feeding stylet and copulatory spicules are formed from
cuticle.
b) Hypodermis- The hypodermis is cellular or partially cellular layer. It lies beneath
the cuticle and contains longitudinal thickening between the somatic muscles that
contain the nuclei, mitochondria, lipid droplets, endoplasmic reticulum,
longitudinal nerves and the canals of the secretory – excretory system. Most
nematodes have four hypodermal chords (one dorsal, one ventral and two lateral
chords). Contains hypodermal glands
c) Muscle layer- The somatic muscle cells are arranged in a single layer. The muscle
cells are spindle shaped and attached to the hypodermis throughout their length. A
non-striated, non-contractile portion of the muscle cell contains the nucleus and
other cell organelles. It is connected to the nervous system by an elongated process
of non-contractile portion of the muscle cell. The stimulation of muscles by the
dorsal and ventral nerves causes contractions in the dorso-ventral plane and results
in the characteristic sinusoidal movement of the nematode.
2. Inner body tube:
The digestive system of nematodes includes the stoma, oesophagus, intestine
and anus or posterior gut. The inner body tube is divided into 3 main regions.
a) Stomodeum: which constitutes the stoma, oesophagus and cardia
b) Mesenteron: which constitutes the intestine
c) Proctodeum: which is the posterior –most region comprising rectum and
anal opening.
a) Stomodeum- Stoma is the portion of the inner body tube lying between the oral
opening and oesophagus. The stomatal opening is small and slit like and surrounded
by six lips with 16 papillae. Two sub-dorsal, two sub-ventral and two lateral. Plant
 parasitic nematodes are armed with a protrusible stylet which is usually hallowed
and functions like a hypodermic needle.
b) Mesenteron- The nematode intestine is a simple, hallow, straight tube consisting of
a single layer of epithelial cells. The intestine is generally divided into 3 regions
viz., 1. anterior or ventricular region, 2. the mid-intestinal region and 3. The
posterior or pre-rectal region.
c) Proctodeum- This comprises rectum and anus. The intestinal tube is connected
with a narrow small tube at the posterior end, through a valve known as rectum. It
regulates the flow of undigested food material which is to be passed outside the
nematode body through a ventrally located aperture known as anus. In male
nematode, the rectum joins with the hind part of the testis forming a common
opening known as cloaca. In female, there is a separate opening.

3. Body cavity or pseudocoelome:

A pseudocoel is a kind of body cavity extending from the mouth to the anus
between the muscle sheath and internal organs found in the nematodes which has no
specialized living cells like epithelium other than those tissue covering the organs
surrounding it unlike that of a true (coelom) which has a mesodermal lining.
a) Reproductive system:
The males are generally slightly smaller than females. The nematodes are
dioecious or amphigonus having a separate male and female within a species.
Generally, the males are lesser in number than females or even be completely absent.
This indicates a tendency towards hermaphroditism and parthenogenesis. Majority of
plant parasitic nematodes do not exhibit any differences as far as body shape. Both
sexes are vermiform.
Female Reproductive system:
The female reproductive system comprises a tube at the top of which the tubular
ovary is located. The foremost part of the ovary, where production of ova takes place is
known as growth zone followed by maturation zone where further development of eggs
takes place. It is followed by a pouch-like structure known as spermatheca where
sperms are stored or retained. It is followed by a muscular structure known as
quadricolumella which is followed by comparatively broader muscular structure known
as uterus. The uterus opens outside to a ventrally located vulval opening through a tube
known as vagina, which is a cuticularised structure.
• Monodelphic- The nematodes may have a single ovary the female is called as
monodelphic.
• Didelphic- The nematodes may have two ovaries then the female is called as didelphic.
• Prodelphic- When a single gonad is present, it may be either directed towards anterior
to vulva then female is called as prodelphic.
• Opisthodelphic- The gonad either directed towards posterior to vulva then female is
opisthodelphic.
• Amphidelphic- The two ovaries are opposite to one another, such as one is anteriorly
directed and other posteriorly directed.
Male Reproductive system:
The production of sperms takes place in testis. In nematodes, whenever the
number of testis is one, it is known as monorchic conditions and when they are two in
number, the condition is known diorchic. The cloacal region is followed by a
protrusible spicule which moves forward and backward with help of specialized
muscles attached with its head region. Spicule is narrower at its tip. A cuticularised
structure lying beneath the pair of spicule is known as gubernaculum which helps and
gives support in movement of the spicule. At the tail end, two filamentous cuticular
expansions are found and they are known as bursa or caudal alae. The bursa helps to
hold the female during copulation.
b) Nervous system:
In plant parasitic nematodes, the nervous system is not well developed. Though
they possess very primitive type of nervous system, they also respond to different
stimuli. The nervous system in plant parasitic nematodes is of two types
1. Peripheral Nervous System
2. Central Nervous System
1. Peripheral Nervous System:
It is located in the periphery which mainly includes body cuticle and also the
cephalic and caudal regions. The parts of nervous system located are well connected
with the nerve ring (circum oesophageal commissure) which encircles the isthmus
region of oesophagus and are considered to be the most important part of the nervous
system (brain of nematode). The peripheral nervous system includes the sensory organs
such as cephalic papillae, amphids, cephalids, hemizonid, hemizonions, deirids,
phasmids and caudalids.
2. Central Nervous System:
Central nervous system comprises the nerve ring which is a band of nerve fibres.
It encircles isthmus. Nerve ring governs various functions of the nematode body. Nerve
ring consists of dorsal gangalion, ventral gangalion, lateral gangalions from which
nerves extending throughout the body. The dorsal nerve is said to be chiefly motor and
lateral nerves mainly sensory in function. The sub – median and the ventral nerves are
partly motor and partly sensory.
c) Excretory system:
The excretory system is not well developed in nematodes. The excretory pore
is located in the anterior mid-ventral line close to the nerve ring. The position of
excretory pore may vary in different genera and even in different stages of the same
species. The excretory systems in nematodes are of two types
1. Glandular type
 2.Tubular type
1.Glandular type: The glandular type consists of a single specialized cell known as
renette cell. It has a posteriorly located enlarged gland known as excretory gland or
ventral gland. This gland is connected to the excretory pore by a duct that terminates in
a pouch like structure known as ampulla. This type is found in members of the class
Adenophorea.
2. Tubular type: The tubular type of excretory system consists of four – cuticularised
canals. Two are anterior and another two are posterior canals. There is a pouch like
structure in the middle which connects both the lateral canals. It is known as excretory
pore.
 TAXONOMY OF NEMATODE
Group: Invertebrata; Kingdom: Animalia

Phylum: Nematoda

Class: Class:
Adenophorea Secernentea

Order : Dorylaimida Order: Tylenchida

Sub Order: Sub Order:

F:longidoridae , G: Tylenchina Aphelenchina
Longidorus ,
Xiphinema
F: Aphelenchoidae , G:
Super Family: Tylenchoidea Super Family:Criconematoidea Aphelenchoides
F: Trichodoridae , G: Trichodorus
, Paratrichodorus F:Anguinidae G: Anguina , F: Criconematidae , G:
Ditylenchus Criconemella ,
F: Delonolaimidae , G: Hemicycliophora
Beionolaimus , Tylenchorhynchus

F: Pratylenchidae , G : F: Tylenchulidae, G:
Pratylenchus , Radopholus Tylenchulus

F:Paratylenchida ,
G:Paratylenchus

F: Hoplolaimidae , G: Hoplolaimus ,
Rotylenchus , Helicotylenchulus ,
Rotylenchulus

F: Heteroderidae G : Globodera ,
Heterodera , Meloidogyne
 CLASSIFICATION OF NEMATODES
The nematode feeding on plants can be divided into above-ground feeders and below-
ground feeders.
I. Above-ground feeders
A) Feeding on flower buds, leaves and bulbs
i. Seed gall nematode – Anguina tritici
ii. Leaf and bulb nematode – Aphelenchoides
iii. Stem and bulb nematode –Ditylenchus
B) Feeding on tree trunk
i. Red ring nematode – Bursaphelenchus cocophilus
ii. Pine wilt nematode – Bursaphelenchus xylophilus
II. Below ground feeders
A) Ecto parasitic nematodes:
These nematodes live freely in the soil and move closely or on the root surface, feeding
intermittently on the epidermis and root hairs near the root tip.
i. Migratory ectoparasite: These nematodes spend their entire life cycle free in the soil,
feeding externally on the host plants, and depositing eggs in the soil. When the roots are
disturbed, they detach themselves.
E.g. Criconemoides spp., Paratylenchus spp. and Trichodorus sp.
ii. Sedentary ectoparasites: In this type of parasitism the attachment of nematode to the root
system is permanent but for this, it is similar to the previous one.
E.g. Hemicycliphora arenaria and Cacopaurus pestis.
B) Semi-endoparasitic nematodes:
The anterior part of the nematode, head and neck are permanently fixed in the cortex
and the posterior part extends free into the soil.
E.g. Rotylenchulus reniformis and Tylenchulus semipenetrans.
C) Endoparasitic nematodes:
The entire nematode is found inside the root and the major portion of the nematode
body is found inside the plant tissue.
i.Migratory endoparasite: These nematodes move in the cortical parenchyma of the host root.
While migrating they feed on cells, multiply and cause necrotic lesions.
E.g. Hirschmanniella spp., Pratylenchus spp and Radopholus similis
 ii. Sedentary endoparasite: The second stage juveniles (J2) penetrate the rootlets and become
sedentary throughout the life cycle, inside the root cortex.
E.g. Heterodera spp. and Meloidogyne spp.
 BIOLOGY OF NEMATODES
The life cycle of a nematode has six stages. The egg stage, first-stage juvenile (J1),
second-stage juvenile (J2), third-stage juvenile (J3), fourth-stage juvenile (J4) and the adult
stage.
The Egg:
The nematode eggs are oval in shape. The eggs are covered by three membranes, 1. the
external protein layer which is the secretion of uterus wall, 2. The middle chintinous layer or
true shell secreted by egg itself and 3. Inner lipid layer (vitalline layer). The chitin content in
the egg shell varies in different species of nematodes. The first moult take place within the egg
and J2 ruptures the egg shell and hatch out. Before hatching, the J1 can be seen wriggling inside
the egg shell.
Hatching (Eclosion):
The emergence of the juvenile from egg is called as eclosion or hatching. It occur either
in response of stimulus or stimuli from the host or takes place under normal environment
condition. The eggs of Globodera rostochiensis generally hatch in response to root exudates
by host plant. The juveniles make a series of thrust with help of stylet on the egg shell @ 40-
90 per minute and finally juvenile emerge out by breaking the egg shell at perforated places.
Moulting (Ecdysis):
The juvenile cuticle is shed after each moult is called as ecdysis or moulting. The
hatched juvenile resembles the adult except for the body size and gonad development. Growth
in nematodes is associated with moulting which occur four times and there are five stages.
After 4th moult the nematodes becomes fully grown adult. During the moulting the entire
cuticle will shed. The Plant parasitic nematode fixes its feeding site in different region of the
root.
Feeding:
Being obligate parasites the juvenile on hatching must find a suitable host for feeding.
Once the feeding site is selected, the juvenile inserts its needle like stylet into the host cell. The
enzymatic secretions from dorsal oesophageal glands start flowing through the lumen of the
stylet into the cytoplasm of the host cell where extra corporeal digestion takes place. At this
stage the highly muscular median bulb starts pulsating at a very fast rate. As the muscles
contract, the lumen of the median bulb is dilated, thus creating a negative pressure that sucks
the semi-digested cell cytoplasm into the nematode stoma. As the food passes through the
median bulb, the secretions from sub-ventral oesophageal glands mix with the food, which is
pushed into the intestine for further digestion, assimilation and absorption. Oesophago-
intestinal valve (cardia) located at the junction of oesophagus and intestine ensures the
unidirectional flow of food. As the cell sap of the host cell is exhausted, nematode withdraws
its stylet and moves to a new site for further feeding. The sedentary endoparasites and semi-
endoparasites which do not move after settling at one feeding site ensure uninterrupted flow of
nutrients for themselves by inducing certain adaptive cellular changes in the host tissue.
Life cycle:

After feeding is over, second and third moults generally occur in quick succession
giving rise to third and fourth stage juveniles (J3 and J4 respectively). Generally the juvenile
forms resemble their adults except in size and development of reproductive system. The size
of the nematode increases at every successive moult. After fourth and final moult, the adult
male or female are formed. Though the precursors of reproductive system start showing in J2,
completely developed reproductive system with vulval opening (in female) and spicules,
gubernaculum and bursa (in male) appears only in adults. The duration of life cycle differs
from species to species and is largely influenced by environmental factors like temperature,
moisture, soil type, aeration and availability of host plant. The normal duration of egg-to-egg
life cycle of most of the plant parasitic tylenchids under sub-tropical and tropical conditions is
25-30 days. Aphelenchids have a short life cycle of 7-10 days. Some dorylaimids complete one
life cycle in 1-2 years.
Reproduction:
Nematodes are basically dioecious, (separate males and females) and amphimictic
(sexually reproducing) which need to find their mates for survival. The sex attractants
(pheromones) released by the females are perceived by the phasmids of the males for mating.
During mating, the protrusible spicules are inserted into the vagina of females. Bursa, the wing
like cuticular extension in the tail region of male nematodes supports it to clasp the female
during mating. Sperms pass through the cloaca into the female genital tract and are stored there
in the spermatheca. As the oocytes pass through the spermatheca, the sperms fertilize them.
Additional layers of the eggshell are formed in the uterus before the eggs are laid by females.
Reproduction by Parthenogenesis (Gk. Parthenos=virgin+ genesis=come in to being) is often
observed in nematodes species in which males are either rare or absent. In such cases young
ones develop from unfertilized eggs. Parthenogenesis is commonly observed in Meloidogyne,
Heterodera etc. Hermaphrodites (Herma= male Aphrodite= female) in which gonads are
capable of producing both male and female gametes for self-fertilization are extremely rare in
nematodes.
Oviposition:
The oviposition behavior in nematodes varies depending upon their mode of parasitism.
Most of the soil dwelling ecto-parasites lay eggs singly in soil itself. Migratory endo-parasites
generally lay the eggs singly inside the plant tissue. Sedentary endo-parasites lay their eggs
clumped together in a gelatinous substance (egg mass). While rectal glands secrete gelatinous
matrix in root knot nematode, in citrus nematode it is the secretion of excretory system. In cyst
nematodes the eggs are retained within the body of mature female. The fecundity (number of
eggs laid by a female) also differs in different nematodes. Generally, it ranges between 10-30
for ectoparasites and migratory endoparasites and 50-500 in sedentary semi-endoparasites/
endoparasites.