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SOME OBSERVATIONS ON THE NERVOUS SYSTEM OF A BUTTERFLY
DISSERTATION SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS
FOR THE AWARD OF THE DEGREE OF
jlagter of ^^ilogopfip IN
ZOOLOGY IN THE FACULTY OF LIFE SCIENCES
THE ALIGARH MUSLIM UNIVERSITY, ALIGARH
BY
MASOOM ZEHRA
DEPARTIVIENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY
ALIGARH (INDIA)
1 9 8 9
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O'
ALIGARH MUSLIM UNIVERSITY
Humayun Murad Ph. D
HEADER
D E P A R T M E N T OF Z O O L O G Y
ALIGARH - 202 002, INDIA
May 21, 1990
This i s to c e r t i f y tha t the d i s s e r t a t i o n
for MoPhil. degree in Zoology of the Aligarh Muslim
Univers i ty , Aligaxh has been completed by Miss Masoom
Zehra, under ray supejrvisiono I t i s o r ig ina l in nature
and I have permit ted the candidate to submit i t fcr
the award of M.Phil, degree in p a r t i a l fulf i lment of
the MoPhilo requirements in Zoology,
( Humayun^ Murad ) Suriervisor
ACKNOWLEDGEMENTS
The author wishes to express her most sincere appre
ciation to Dr, Humayun Murad for his precious guidance, contin
uous encouragement, critical and constructive suggestions
during the progress of research without which this work would
never have come to light. I am deeply indebted to him indeed.
I feel highly obliged to Prof. Mumtaz A. Khan,
Chairman, Department of Zoology for providing necessary
laboratory facilities,
I am very much thankful to Kr, Ammar Ahmad Khan for hel
ing me financially during the course of my studies.
It is my pleasant duty to acknowledge the assistance
of my lab. colleagues Dr. Muzaffar Atiqui, Dr. (Miss) Archana
Bansal, Miss Robina Naseer and Mr. Mohd, Amir, whose help
proved to be very useful during this research work .
I have no words to express fully the indebtedness,
benevolance and gratitude I owe to my parents.
\ % ^
MASOOM 2EHRA
C O N T E N T S
Page No«
INTRODUCTION 1 - 8
REVIEW OF LITERATURE 9 -15
MATERIAL AND METHODS 16 -17
RESULTS AND DISCUSSION 18 -27
REFERENCES 28 -44
•k-kieitleiiititit
INTRODUCTION
The Monarch butterfly Danaus chirvslppus (L) belongs
to the order Lepidoptera and family Nymphalldae which is a
dominant family among lepidopterans. The characters of this
family are; forelegs in both sexes are reduced in size usually
folded under the thorax and functionally impotent, the tibiae
are short and clothed with long hairs hence the name "brush
footed butterflies" Imms (1957)» It is usually seen during the
months of May and June, They are found in India, New Zealand
and Australia. It feeds on milkweed Asclepias curassavica but
but had now been observed feeding on Gossvpium arborium
(musturd). These Lepidopterans are one of the most familiar
and easily recognizable among insects and in this order
coloration has reached the highest degree of specialization.
These insects have always been popular objects to study.
Economically, Lepidopterans are of great importance in their
larval stages, as they bear cutting type of mouthparts.
Majority of injurious species devour the foliage and shoots
of trees and crops; a smaller number bore into stem or attack
underground parts. Several species are injurious to timber,
others attack manufactured goods as carpets, clothings and like
wise, some are extremely destructive to stored grains. Some of
these insects create great trouble to agriculturists and
farmers.
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According to Roonwal (193 6, 193 7) the central nervous
system originated as a thickening of the ectoderm on either
sides of the mldventral line. At about the time of segmentation
and prior to the dorsal closure of the blastoderm to form an
embryo rows and columns of large neuroblasts arises from beneath
the superficial cells of the ventral epidermis.
Considerable work has been done on the nervous system
among different insect groups as Hymenoptera, Diptera and
Coleoptera but little work has been done about the nervous
system of lepidopterous insects. The present study is concerned
with the Central Nervous System of a Monarch butterfly D^naus
chrvsippus (L.). A generalized picture of the lepidopteran
nervous system was given by Colhoun (1935), Treherne (1966)
and House (1977). Chauthani and Callahan (1967), Kuwana (1932)
described the nervous system of silkworm Bombvx mori. In 1961
Ehrlich and Davidson successfully studied the internal anatomy
of Danaus Plexippus (L) (Nymphalidae).
The nervous system of a butterfly is divided into three
divisions, the Central nervous system, the Peripheral nervous
system and the Sympathetic nervous system. The Central nervous
system is a conducting system ensuring rapid functioning and
coordination of effectors, modifying their responses according
to the input of peripheral sense organs, Snodgrass (193 5) studied
the evolution of the annelid - arthropod nervous system from
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whlch It is very clear that the first central group of nerve
cells had its origin in the ectodeirtn at the anterior pole of
the body forming a small ganglion associated with a sensory
apical plate. Later there appeared several groups of paired
sensory cells at the bases of tentacular or other sensory organs
behind the apical plate. The central neuroendocrine system of
insects control many aspects of physiology and behaviour,
Raabe (1982), Truman and Taghert (1983) studied the
endocrine organs in detail. Cell and tissue culture have made
a major contribution to our understanding of biochemistry,
physiology and morphology of invertebrate nervous system.
Nervous system basically consists of brain which is principal
association centre of the body receiving sensory input from the
sense organs of head and via ascending intemuncial fibers from
the most posterior ganglia. The brain of most lepidopterans
species has been studied in a generalized pattern by
Bretschneider (1921), (1924 a,b), Hanstrom (1925), Schrader
(1938), and Ehnbom (1948), Others like Kenyon (1896),
Bretschneider (1913), Hanstrom (1928), Porer (1943), Bullock
and Horridge (1965) gave description of adult lepidopterans
brain microanatomy. Brain, as it is the computer or the master
organ of the body regulates different vital functions as growth,
reproduction metamorphosis etc. Our present understanding of
the function of the nervous system provides an excellent example
of the cumulative values of the complementary approaches to a
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single problem, ultimately the function of nervous system
becomes the province of the behaviourist concerned with the
Ways in which the animal responds to the environment. The
Central nervous system of Danaus involves a marked expansion
of the brain and outward rotation of the protocerebral lobe
and a fusion of sub-oesophageal ganglion with a ventral tri-
tocerebrum. The attention of insect neurophysiologists is no>
being focussed on the central nervous system morphology and
pharmacology with special reference to generation of behaviour.
The primitive brain lies above the anterior end of oesophagus.
Brain structure and behaviour was studied by House (1977)
Smith (1962) pointed out that a shortage of time and patience
on the part of the morphologist is the chief reason for the
lag in providing an account of the insect nervous system. Th-:̂
available literature reveals that majority of the researchers
on this very particular subject, especially in Lepidoptera
have confined themselves to the larval forms as they are
destructive in this very stage. Du Pbrte (l9l5), Swaine (1920),
Hillemann (1933), Bickley (1942), Likky (1959), Srivastava
(1959), Mathur (1967), Hanstrom (1925), Kuwana (1932), Nuesch
(1957) have contributed to the knowledge of nervous system in
adult lepidopterans, but their studies were restricted to the
nerve supply of one or more organ system. Structurally Insect
brain is a compact creamish bilobed stiructure and is continuous
laterally with expanded optic lobes, it consists of various
parts as first brain or Protocerebrum which bears optic lobes.
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deutocerebrum or second brain bears antennal lobes which is
divided into sensory and visceral motor areas, and thiirdly
tritocerebrum which is the smallest portion of the insect brain
containing a pair of lobes beneath the deutocerebrum of the
ventral nerve chain of segmental ganglia; the first is suboeso-
phageal ganglia, which is formed by the incorporation of the
gnathal segments, suboesophageal ganglion lies in the ventral
region of the head beneath the oesophagus, thoracic and abdominal
ganglion are all connected by longitudinal nerve cord or double
connectives to form the ventral chain, Suboesophageal ganglia
is a compound structure lying ventrally in the head region,
arising from the fusion of mandibular maxillary and labial
segments.
Typically there are three thoracic ganglia each with
five or more nerves. The number of abdominal ganglion occuring
in lepidopterans are four and the last one is always compound
being derived from the ganglia of the last four abdominal
segments.
In majority of insects some degree of fusion particularly
in the abdominal ganglion, which is smaller than the thoracic
ganglia connected across the midline (Mathur 1969, Wigglesworth
I960, Welnbormair, Pohlhanmer, Dumberger 1977) and joined with
the ganglia of adjacent segment by a pair of connectives known
as intergangllonic connectives which have no cell bodies. Basic
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elenuants of nervous system i s neurone, derived in the course
of development. Neurones are of several types v i z . Bipolar
and Multipolar (Wigglesworth i960) . Most of the i n s e c t s have
monopolar neurone. I t has a nucleated c e l l body and a long
cytoplasmic project ions which extends to make contact with
other neurone. The ce l l body i s perikaryon while project ions
a re known as axon. The un i t of Insec t neurone system transmit
e l e c t r i c a l disturbances or impulses from one part of the body
to the o ther . They do not occur s ingly but are conjugated as
proved by Gilbert (1973), Brain as we a l l know regula tes body
metabolism, digest ion excret ion, metamorphosis, growth, repro
duction, feeding, mating e t c . Nervous system exhib i t s cer ta in
amount of concentration with regard to the ganglia of the ventral
nerve cord.
The most extensive endocrinological s tudies was made by
William and his co-workers on the diapausing pupae of Gecropia
silkworm. In general the endocrine system of insec t consis ts
of (as described by Wigglesworth (1972) in his recent volumes)
i s ( i ) modified nerve c e l l s , neurosecretory c e l l s ( i i ) neuro-
haemal organs as corpora cardiaca and corpora a l l a t a , and
( l i i ) independent organs of i n t e rna l sec re t ion . Here i t i s
necessary to know about the neurosecretion and how neurosecretory
and ce l l regula tes d i f fe ren t hormonal balances. The term neuro
secre t ion means d i f fe ren t things to d i f fe ren t peoples, and these
c e l l s are able to sus ta in an effect on t h e i r t a rge t organs
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(Hlghman I96I, Delphin 1963). According to Gilbert: and Kerkut
(1985) in any one neurosecretory cells the secretory product is
accumulated in membrane bound vesicle which are usually though
not in all cases of rather uniform size and appearance. Neuro
secretory cells has been studied in various insects as fine
structure of these organs was described in Rhodnius prolixus
by Kuster and Davey (1981). The allotropic activity of median
neurosecretory cells of GaUeriq mellonella^ a wax moth was
successfully observed by Muszynska - Pytel (1986). He described
that the brain allotropic hormone (ATTH) is released in Gallarla
mellonella from the median neurosecretory cells present in pars
intercerebralis. Furthermore in vitrx) neurx)secretion can be
used as a probe for locating and exploring neurohaemal sites,
measuring reasonable stores of neurohormones studying excitation,
secretion, coupling and obtaining the circulating forms of
neurohormones as said by Highnam (1961), Delphin (1963),
Treheme (I966) experimentally proved that some neurones in
insect ganglia are neurosecretory i.e. the nerve cell bodies
are modified to produce quantities of granules called elementary
granules or neurosecretory droplets. After synthesis and
condensation into granules within the neuronal perikarya the
neurosecretory material is apparently transported to neurohaemal
organs or storage via the axons. Usually nerve cells convey
uniform action by releasing specialized chemical substance
which affects other cells, neurone system is modified in distinct
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ways and provides three attributes, i.e. of specificity, speed
of delivery and length of message. The release of neurosecretory
products from the membrane bound vesicles in the cytoplasm to
the outside of cell which in turn affects the target organs and
stimulate it to release the hormone essential for many vital
activities of the body, Neurohaemal organs such as corpora
cardiaca and corpora allata which are paired in some insects
but in this particular species they are fused with each other
in the cervix region. The corpora cardiaca and oesophageal
ganglion govern the function of the dorsal vessel. These are
anteriorly connected, with the protocerebrum through a pair of
nerve called procardiac nerves. The corpora allata are very
small lying ventral to and in very close association in fusion
state with corpora cardiaca with which it is connected by means
of a very short connecting duct which are not normally noticed.
The present study has been conducted with a view to
provide adequate information on the morphology and histology of
the insect central nervous system with particular reference to
REVIEW OF LITERATURE
Description of adult insect brain microanatomy still
needs some clarification. Some of the authors such as Kenyon
(1896), Bretschneider (1913, 1921, 1924 ab), Henstrom (1928),
Power (1943), BullocX and Horrldge (1965) gave a detailed review
about the anatomy and morphology of an insect brain, Kenyon
(1896), Cajal and Sanchez (1921); Schrader (1938) gave only
some details on particular type of cells which are present in
the brain region while others as Bretschneider (1914 ab),
Holste (19 23); Alverdes (1924), Jawlowsld. (1936), Bierbrodt
(1924), Rogoff (1943) have studied the morphological differences
between the mature and immature nervous system of insects.
Hanstrom (1925), Hertwick (1931), Schrader (1938), Pyle (1941)
noticed larger size of an adult brain which was due to the
increase in the number of cells and larger and more complex
n euro pile.
Brain anatomy of Monarch butterfly Danaus chrvsippus (L.}
(Nymphalidae) was studied carefully by Ehrlich and Davidson
(1961). Brain of several lepidopterans have been studied in a
generalized pattern - by Bretschneider in (1921, 1924a, 1924b),
Ehnbom (1948) distinguished the following 4 types of lepidopterous
brain: as followsi-
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a) Hesperoidea and Rhopalocera, which have rounded opt ic lobes;
(b) the Macrofrenate families^ except Diepanidae and Cymatopho-
nidae (Weber 1933) which have elongated, l a t e r a l l y projecting
opt ic lobes and in which the deutocerebrum, especia l ly in
Noctuidae, i s r e l a t i v e l y large* (c) Drepanidae» Cymatophonidae,
Pyraloidea, Hepialidae and Tineioidea; which have l a t e r a l l y
compressed brain with op t ic lobes extended dorsoventra l ly
(d) Micropterygidae which have a l a rge oesophageal canal with
r e l a t i v e l y long connective and a pr imi t ive brain s imilar to tha t
of Trichoptere.
The brain of insec t va r i e s i n s i z e from t i n i e r pinpoint
to several cubic mi l l imeters . I t i s not possible to de t a i l the
novel arrangements met wilhin d i f fe ren t species because of
lacunae in our knowledge. The a t l a s of i n sec t brain prepared
by Straflsfeld (1976b), i s a landmark in the study of b ra in .
According to Vil lanes (1884) the brain of lepidopterous
i n s e c t s i s divided in the th ree pa i r s of neuromeres designated
as protocerebrum, the f i r s t b ra in , deutocerebrum, the second
brain and l a s t l y the t r i tocerebrum, the t h i rd bra in , which i s
the minutest port ion among the t h r e e .
Deta i l s on the Coleopteran bra ins are only few but
Abraham in (1969) gave a very i l l u s i v e account of the histology
of the bra ins in Dvtiscus maroinal is .
- 1 1 -
Murrary and Tlegs (1935) described the histology of an
insec t b ra in , Sa t i ja and Dass (1963) gave a deta i led account
on the s t ruc tu re of the bra in of a b e e t l e Onthophaous c a t t a .
The next region of the cen t ra l nervous system i s the thoracic
region which i s formed by the fusion of prothorax, mesothorax
and metathorax. The lepidopterans bear thorac ic nerve centers
in the mesothoracic port ion of the thorax region as i l l u s t r a t e d
by Hachlow (1931); Similar conclusions were drawn by Pflugfelder
(1958) but they did not find any appropriate endocrine glands
in the mesothorax. Pakuda (1940) observed t h a t the t ransplan ta
t ion of the gland located near the prothoracic sp i r ac l e could
s t imulate pupation in the i so la ted hind par t s of Bombvx larvae .
This a t t r ac ted the i n t e r e s t of insec t endocrinologists to the
prothoracic glands,
Prothoracic gland sjTithesize ecdysone or substances
which have ecdysone l ike a c t i v i t y which was l a t e r confirmed by
Takeda (1976), The physiological p roper t i es of ecdysone have
been studied careful ly by Novak (1966); Sorm and Slama (1974),
which waxes and wanes under the d i r e c t control of cen t ra l
nervous systan which in t imate ly re leases a t rophic hormone
general ly known as prothoracico- t rophic hormone or bra in hormone,
and as described by Fakuda (1944), Williams (1947, 1952) i t
regresses or depresses the terminal moult on the diapause break.
According to Takeda (1977b) in Monema flavescens the neuro
secre tory substance, the prothoracicotrophic hormone are
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released from neurosecretory B type of c e l l s present in the
construct ion of the brain known as pars i n t e r c e r e b r a l i s .
Histology of prothoracic glands had been studied by
Herman (1957). The most appropriate work in the f ie ld of
neurology about the thoracic portion of the insec t Central
nervous system was done by Williams and his co-workers (1947)
on diapausing pupae of Cecropia silkworm. After these in for
mative discover ies they had come to the conclusion tha t the
imperative ro le of the bra in hormone i s very necessary for the
regula t ion of metamorphosis which was l a t e r on confirmed by
Williams (1946, 1952).
More recent ly other researchers have proved t h a t
prothoracic glands in the silkworm, Bomby?̂ mori (Chino, Sakurai,
Ohtaki, Ikekawa, Miyazaki, Ishibashi and Al»uki, 1974), the
tobacco hornworm, Manduca ^e^ta (King, Bollenbacher, Borst,
Vedeckis, 0 ' connor, I t tyche r i ah and Gi lber t , 1974), the cockroach,
Leucophaea maderae (Borst and Engelmann 1974; King and Marks
(1974) and the meahworm, Tanebrlo molitay produce ^^-ecdysone.
Srivastava and Singh (1973); and Hintz Poodufal (1987)
studied about the neurohaemal organs in lepidopterans and they
suggested the t ranspor t of neurosecretory granules to the gland
c e l l . On the bas is of experimentation by Srivastava, Tiwari
and Kumar (1976) i t may be assumed tha t the innervat ions of
prothoracic glands are e i t h e r involved in feeding, sensory
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proprioceptive inputs to the Central nervous system or exert
a nervous inhibition on the activity, (Synthesis or release of
ecdysone) of the prothoracic gland. The whole issue of the
significance of the prothoracic gland innervation is therefore
investigative. Albrecht (1953) studied the anatomy of the
migratory locust. Locust miaratoria mlaratoides? but later on
the nervous system of Cockroach Periplanata amerlcana was
illustrated by Pipa and Cook (1959); and Shankland (1966),
As the research proceeded, people tried to find out more
details or to explore the nervous system of various insects as
Maki (1936) studied the nervous system of alder fly Chaulioides
formosanus , In so far as lepidopterans are concerned there
have been numerous general discriptions but the detailed study
of Hyalophora larvae described by Libby (1959) proved to be
very useful in neurology. As the work proceeded various other
insects of the same order have been taken into consideration,
the thoracic nervous system of Telea Polvphemus was described
by Niiesch (1957) and Eaton (1974), Eaton (19 74) also described
the anatomy of head and thorax of the adult tobacco horn worm
Manduc^ sexta. Cook (1980) studied the nerve architecture of
the oviduct in Pectinophora aosavpiella. The terminology
used for naming of the nerves is the same as that used for the
study of innervations of head and thorax as described by Eaton
(1974). Libby (1961) discussed and successfully explored the
branches of nerves in the sixth and tenth segment in males of
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Hyalophora, According to Srlvastava, Tlwari and Kumar (1976)
innervation of Prothoracic gland is found in at least five
insect orders, while the function of these nerves have been
variously suggested by Lafon, Gazal, Roussel (1976); Goldsworthy
(1971); Millis Greenslade, Couch (1966); Uroman Kalpanis Robbinc
(1971), Walker Bouley (1971); Vanthan der lea (1965).
The Central nervous system of insects also includes the
abdominal ganglionic masses. Each ganglion which is somewhat
bulb like in shape is connected by interganglionic connectives
as experimentally proved by Siegler and Burrows (l979).
Newport, (183 4); Brandt 1879; Paterson, (1899), Buxton
(1917) exhibits a certain amount of concentration with regard
to the ganglia of the ventral nerve cord. The conditions of
thoracic and abdominal ganglion differ greatly as found in
Hepialus in which three thoracic and five abdominal ganglia
was observed. In Tinea pellipnella (Micropterygidae) and some
other insect spp., the 4th and 5th abdominal ganglia are fused
into a large common centre. The majority of lepidopterans are
characterised by two thoracic and four abdominal ganglia; those
of the raeso- and raetathorax are fused and the abdominal ganglia
lie in the 2nd to 6th abdominal segments. While in Fumea
intermedia and others there are four abdominal ganglia in both
sexes. The terminal ganglion in some insects gives out several
pairs of nerves as shown by Libby (1959) in Jg. cecropja. The
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intemal morphology of lepidopterans described by numerous
workers as in one of the most complete account given by
Osteo and Helms (1976) in Laxaarotls albicost^ belongs to
the family Noctuidae, Ehrilch and Davidson (1961) gave gross
anatomy of Danau;̂ plexippus« which forms the baseline in the
study of lepidopteran nervous system, and proved to be very
successful in the foregoing discussion.
Neurosecretion has recently been the subject of rather
great interest and prominent work in the field of neuro
secretion was done by Van der Kloot (I960), Gane (1966). The
term neurosecretion was defined differently by different peoples.
According to Van der Kloot (i960). Neurosecretory cells are
those nerve cells which when examined microscopically shows
signs of secretion. Some of these cells are involved in the
release of hoirmones, while other release substance which
effects only for short duration. Some neurones in insect
ganglia are neurosecretory i.e. the nerve cell bodies are
modified to produce neurosecretory droplets. Most of the work
on neurosecretion is done electron microscopically. Some evi
dences reveal that the ganglia of the ventral nerve cord
contains many neurosecretory cell, (Nayar 1954, Wigglesworth
1955a, Fraser 1959, Geldiay 1959). In insect number of different
neurosecretory cells has been described.
MATERIALS AND METHODS
Eggs of D^naus chrvs lppus (L) (Monarch b u t t e r f l y ) were
col lected from plants of AsclepJas curraasavlca (Akawwa) from
t h e Universi ty campus and Aligarh Port , and rear ing was done
under laboratory condi t ions .
Adults were kept i n a cage in a basement laboratory
receiving daylight but never d i r e c t sunl ight . Suppl«nentary
i l luminat ion was derived from 60W bulb placed away from one
side of the cage and run overnight during winter season. The
temperature during night was e s sen t i a l for reasonable fecundity
and adult l i fespan . Adults were fed on cotton soaked sugar
solut ion and adult l i fespan was about 20-22 days with the
temperature 29 + I 'C ,
Materials were drawn from the maintained standard
cu l t u r e .
For morphological and anatomical observations the
i n sec t s were anaesthesized and were dissected in Normal Saline
under the binocular microscope. Methylene blue proved to be
very sa t i s f ac to ry in d is t inguish ing the ganglion and i t s
innervat ion. For h i s to log ica l s tud ies material was taken and
dissected in physiological s a l ine and then kept in Bouins fluid
(alcoholic) for about 12 h r s ; dehydration was done with different
-17-
alcohol ic grades. Material was cleared in xylene and absolute
alcohol/ with a r a t i o of 50i50 and f i n a l l y t ransferred to pure
xylene used as c lear ing agent. The claved material was t r a n s
formed t o paraffin wax of 50-60 m.p. for about 4 h rs . Blocks
were cut at 3 xi with a rocking microtime.
For s ta ining purposes Heidenhain's i ron haematoxylin
and alcoholic Eosin were used for cytoplasmic and nuclear
d i f f e r e n t i a t i o n . Complete dehydration was ensured before
mounting of the material in D.P.X, mountant for microscopic
examination.
Results and Discussion
The insect Danaus chrysippus (L) undergoes following
biological stages to complete its life cycle,
(1) :^3G STAGEt The egg is generally laid on the lov;er
surface of the leaf; It may also be laid, more rarely on the
upper surface. It is generally glued to the surface near the
margin of the leaf on one side of the midrib. It is creamy
white nearly globular, slightly longer along its longitudinal
apex. The base is round and flat and each egg is laid singly
and one egg on one leaf only, as a irule, but in one case seven
eggs were found on one leaf. Big or small leaves are indiscri
minately choosen for egg-laying. The surface of the egg is
marked by ribs running from the basal periphery and converging
into the apex of the egg. Between these longitudinal ridges
there are faint parallel striations situated one below the
other. The longitudinal ridges do not meet each other at the
apex but leave a micropylar area reticulated. The egg is set
vertical on leaf surface,
THE NSWLY HATCHED LARVAt The larvae on hatching from the eggs
at once attack the empty shells and in nearly all cases devour
them to well near the bases begining from the apex and after
words proceed to feed on tender leaves by the margin. It is
cylindrical in shape almost uniform in dimension throughout.
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t a p e r i n g v e r y s l i g h t l y p o s t e r i o r l y . The head i s l a r g e r t h a n
t h e p r o t h o r a c i c s e g m e n t , r o u n d i s h d e f l e x e d . S u r f a c e i s s h i n i n g ^
d o r s a l s u r f a c e h a v i n g an i n v e r t e d Y- shaped s u t u r e . The t h o r a c i .
l e g s a r e s h i n i n g , w e l l d e v e l o p e d d i s t i n c t l y j o i n t e d and g r e y i s h
b l a c k i s h c o l o u r e d . The c o l o r o f t h e body i n c l u d i n g t h o r a c i c
r e g i o n i s n c r e a m y w h i t e , a n a l c l a s p e r s have h o o k l e t s . I n t h e
f i r s t i n s t a r l a r v a t h e s i z e i n c r e a s e s , o n l y c r i m s o n i s h t r a n s -
v e r s b a n d s have d e v e l o p e d from s i d e t o s i d e on each segment .
V e n t r a l c o l o r a t i o n l i g h t y e l l o w i s h , t h a n i n s e c o n d , t h i r d ,
f o u r t h and f i f t h i n s t a r l a r v a e o n l y t h e s i z e i n c r e a s e s .
PUPA; L a r v a e p u p a t e s g e n e r a l l y on t h e f o u r t h o r f i f t h
day by c a s t i n g o f f i t s l a s t m o u l t . The pupa i s suspended
v e r t i c a l l y head downwards from t h e p o i n t o f a t t a c h m e n t by
i t s b l a c k c r e a m a s t e r whose hooked h a i r s a r e e n t a n g l e d i n a
pad of sillf. woven on t h e s u r f a c e , A day a f t e r p u p a t i o n
g o l d e n s p o t i s s een on t h e pupa r ium,
ADULT: I n c u b a t i o n p e r i o d n i s abou t two d a y s . B u t t e r f l y
emerges by b u r s t i n g t h e l o w e r o r head end of t h e pupa and
hangs on t h e empty pupa c a s e w i t h c r i m p l e d wings and s h o r t
t h i c k abdomen. I n a b o u t h a l f an h o u r wings expand and
h a r d e n and abdomen l e n g t h e n s . P e r i o d of l i f e c y c l e i n ho t
w e a t h e r i s 20 d a y s (Egg p e r i o d 3 d a y s , l a i rva^ p e r i o d 10
d a y s , pupafe p e r i o d 7 d a y s ) .
-20-
THE NERVOUS SYSTEM}
The Insect nervous system is divided into (a) central
Nervous system (b) Stomodeal Nervous system and (c) Peripheral
Nervous system.
At the moment the study of the nervous system of
Danaus chrvsippus (L) is only confined to the Central Nervous
System, Here in this foregoing discussion only the anatomy
and morphology of this very system has been taken into
consideration,
THE CENTRAL NERVOUS SYSTEMi
The central nervous system of Monarch butterfly
D. chrvsippus (L) consists of Brain, Suboesophageal ganglion
and the Ventral nerve cord.
The nervous system of D, chrvsippus encircles the
pharynx and persists along the midventral line below the
alimentary tract in the thoracic and abdominal portions. The
present author has observed that in the larvae stage of
monarch butterfly the number of ganglia is according to the
number of body segments but as the stage advances and the
catterpillar moves towards the adult stage the number of
ganglionic masses fuses. In the advance stage a greater
degree of cephalization is observed. The first part of the
insect nervous system is.
- 2 1 -
BRAIN: (Br) Brain l i e s j u s t above the oesophagus between
the supporting apodemes of the tentorium. I t i s somewhat
creamish in color and a compact mass without conspiquous
prominences. Among the chief ear ly wr i t e r s on the s t ruc tu re
of the brain are those of Vialanes (1886-87), Haller (1905),
Janet (1905), Jonessen (1908). Brain occupies major portion
of the cranium. I t l i e s t ransverse to the head capsule. I t
i s the dorsal ganglionic cent re of the head and i s formed by
the coalesence of the f i r s t th ree neuromeres of the embryo.
This t h ree fold d iv is ion i s maintained in the completed organ
which divides in to corresponding regions, which are designated
as Protocerebrum or f i r s t b ra in , Deutocerebrum or second brain
and Tritocerebrum or the th i rd brain respec t ive ly . However
these th ree subdivisions of rthe bra in are not d«narcated
ex te rna l ly but are ac tua l ly d i f fe ren t i a ted on the Igasis of
the nerves or ig ina t ing from than. The Brain of D. chrvslppus
was also studied by Ehrlich and Davidson (1961). Brain i s a
bilobed s t ruc tu re with two bulding ganglia on i t s e i the r
s ide as shovm in Fig. 1. The l a t e r a l port ion of the brain
i s depressed but i t s dorsocaudal region i s s l i g h t l y raised
to form an arch over the stomodeum.
PROTOCEREBRUM (IBr)i Protocerebrum in D. chrvsippus i s
dorsal i n posi t ion and occupies la rge portion of insec t brain
as shown in Fig. 1 and Fig. 15, This comprises the cent ra l
- 2 2 -
complex, pars i n t e r ce reb ra l i s^ corpora pedunculate and
op t ic lobes . I t s two lobes have undergone fusion which i s
v i s i b l e ex terna l ly by a cons t r i c t ion along the mid-longi
tud ina l groove Fig. 1. The cen t ra l complex i s not very
prominent but pars i n t e r c e r e l o r a l i s (Pi) i s very conspicuous
under the l i g h t microscope Fig. 1. I t i s the anteromedian
c e l l u l a r area of the brain occupied by motor c e l l s and
neurosecretory c e l l s . The protocerebrum d o r s o - l a t e r a l l y
gives out a pa i r of s tout nerves going to the l a t e r a l o c e l l i ;
Pi is tological ly the f i r s t brain bears a pa i r of la rge rounded
opt ic lobe (Op.L) and oce l l a r cen t re . The opt ic center
invera tes the eye. Each opt ic lobe i s demarcated from the
protocerebrum along a d i s t i n c t external cons t r i c t ion (Cons.)
Fig. 3. The opt ic lobes according to Chauthani and Callahan
(1967b) in H. zea begin to d i s t ingu ish i t s e l f in the prepupal
s t age . The opt ic ganglion bears lamina, (L) medulla (M) and
lobul la Fig, 4, The neuropiles of the op t ic lobe and the
inne r lobula are conspicuously s t r a t i f i e d and arranged in
columns based upon the ca r t r idges of r e t i n a l elements Fig. 4.
The laminal l in ings are also c l ea r but the medullar portion
i s fake. The most conspicuous port ion of the protocerebrum
are corpora pedunculate (CP) Fig. 1. Each of these s t ruc tures
are semicircular with a s l igh t cup l i k e depression, the
calyx (Cy) and a branching s ta lk (S) the peduncle in the
protocerebral neuir)pile. There are two calyces in each brain
hemisphere.
- 2 3 -
DSUTOGEREBRUM (I I Br) i Behind the Protocerebrum l i e s a deuto-
cerebrum or the second brain which i s vent ra l in pos i t ion .
This i s the middle part of the bra in which i s smaller i n
s ize Fig. 15. I t can however be iden t i f i ed by the antennal
lobe, which bears antennal nerves supplying to the antenna.
In the vent ra l region of the brain the two opposite deuto-
cerebral lobes touch each o ther , and the antennal nerves
a r i se from t h i s very region of the bra in , Abraham (19 67)
reported the presence of a th in and a th ick f ibre system in
the antennal glomeruli. But in Spilachn^ viqintioctomaculata
the antennal lobe lacks the glomerular the fibrous zones.
Bahadur and Srivastava (1968) i n Prodenia l i t u r a have shown
large sized deutocerebrum. The antennal lobes are separated
by some dis tance but t h e i r coramisures l i e within the substance
of the bra in . The antennal lobes have motor and sensory
areas Fig, 15.
TRITQCEREBRUM ( I I I Br): This i s the smallest portion amongst
the t h r ee . I t l i e s ventra l to the antennal lobe from which
i t i s f a in t l y demarcated. These are connected with the pro
tocerebrum on dorsal region of the bra in . I t s pos te ro - l a t e ra l
par t s are drawn downswards to l ink with t he sub-oesophageal
ganglion.
SUBOESOPHAGEAL GANGLIQNt I t i s a compound mass of segregated
ganglia of gnathocephalic segments. The suboesophageal
ganglionic mass lying below the pharynx i s formed by the fusion
-24-
of paired mandibular, maxillary and labial ganglia. Ehrlich
and Davidson (1961) in Q, chrvsippus have recorded absence of
mandibular nerves. Functionally it probably works in close
association with the supra-oesophageal brain (Tyrer and
Gregory (1982), Stransfeld (1976), but morphologically it
resembles thoracic ganglia in nerve distribution Pig, 15,
VENTRAL NERVE CORD (VNC): The ventral nerve cord of
D. chrvsippus bears six dull cream coloured ganglionic masses
arranged in a linear fashion along the mid ventral longitudinal
line of the trunk. Each ganglionic mass is connected by an
interganglionic connectives Fig. 16,
THORACIC G^^GLIQN: In the thoracic portion there present
three ganglionic masses which are designated as Prothoracic
ganglion, mesothoracic ganglion and metathoracic ganglion
respectively.
PROTHORACIC GANGLION (Ptg): It is the first ganglion of the
ventral nerve cord. (1 Gng.). It is smaller than the combined
meso and metathoracic ganglion Fig. 6, Pig, 16 Chauthani and
Callahan (1967a) in H, zeg have shown it somewhat displaced
from its original position in the mesothorax. Anteriorly it
is connected with the suboesophageal ganglion by a fused
pair of ventral nerve connectives.
-25-
Two short but very stout connectives extend
posteriorly from the prothoracic ganglion to connect it
with the second thoracic ganglionic masses. This is the only
place in the nerve chain of the adult where the paired
ganglionic connective maintain their separate identity other
wise they are completely fused with each other. However,
the paired nature of interganglionic connectives is maintained
in the larval stages as shown by Chauthani and Callahan
(1967b). in H, zea. The nerve distribution of prothoracic
ganglion is simply illustrated in Fig, 21. As shown in
Fig. 20, the thoracic ganglion of the ventral nerve cord
bears 5 pairs of nerves and a connective and these nerves
are distributed to various organs.
MESO META THORACIC GANGLIONt These ganglia are somewhat
barrel shaped in structure. This is a composite structure
formed by the fusion of meso-, and metathoracic ganglia
together with first two abdominal ganglia. They are connec
ted with each other and a line of demarcation between the
two is observed very clearly under the light microscope as
revealed in Fig, 11. The cells are very prominent near the
connection Fig, 11. Fig. 12. Anteriorly the mesothoracic
ganglion bears neuromeres Fig. 10. Which are prominent
Fig, 6, While the posterior view of the thoracic ganglion
when seen microscopically, does not show any clear cells.
-26-
The meso- and metathoracic ganglia show ganglionic core
which i s the cen t ra l mass and i t bears ganglionic c e l l s at
i t s periphery. The Central mass has g l i a l c e l l s Fig, 11,
Neuroanatomical s tudies of i n s e c t segmental ganglion
have lagged behind physiological s tud ies . One of the few
organisat ional p r inc ip les f i r s t elaborated by Binet (1894)
r e l a t i n g to the d i s t r i b u t i o n of the motor elements pre
dominantly in the dorsal par t and sensory elements in the
ven t ra l portion of the ganglionic core . This general izat ion
was broadly substant ia ted by the c l a s s i c a l methylene blue
s tud ies by Zawarzin (1924a,b), on the dragon f ly .
ABDOMINAL GANGLIA; (Abd.g,) : The ven t ra l nerve cord in the
abdominal region bears 4 ganglionic masses. These masses
are somewhat c i r c u l a r and creamish in colour and attached with
each other in a chain l i k e manner. Each abdominal ganglion
gives out two pa i rs of l a t e r a l nerves to the segment concerned.
Fig. 15, These nerves innervate v e n t r o - l a t e r a l muscles of the
abdomen and the d iges t ive organs. The f i r s t 3 ganglia are
button shaped and are s i tuated in the pos te r io r halves of
2nd, 3rd and 4th abdominal segments. The nerve d i s t r i bu t i on
of the an te r io r 3 ganglia i s s imi lar Fig. 15, Each ganglion
i s encircled by a membrane known as Neurilemma (Nlm) and has
a ganglionic core (gc) . The c e l l s are seen in c lu s t e r s at
the peripheral region of the ganglion Fig, 14. The terminal
ganglion of the abdominal region of ven t ra l nerve cord i s
-27-
somewhat bulb like in structure. This ganglion is composite
structure made up of 6th, 7th and 8th abdominal segments.
This ganglion differs in size with the other three ganglionic
masses, and this ganglion is complicated in the sense that
it bears more than two pairs of nerves Fig, 16. In males
thus terminal ganglion gives out 5 paired nerves. The first
pair represents the typical segment anterior lateral nerve
innervates dorsal diaphragm, heart segmental muscles and
spiracles. Similarly 2nd pair is typical post lateral nerve
supplying to the muscles. The 3rd pair of nerve arises
posteriorly to the 2nd pair and travel in the postero lateral
direction. The other pairs goes to respective reproductive
organs, Libby (1961) in H, cecropiq. shows ten pairs of
nerves arising from the tenrdnal ganglion, three pairs each
for 6th, 7th and 8th segments and one pair for the reproductive
organs and the genitalia.
^n female there is no differentiation in the inner
vation but only the point of dissimilarity is in the size of
the terminal ganglion. The shape is bulb like and size is
smaller than that of the males. The first pair of nerve is
fairly thick whose branches innervate the sixth segment. Other
pairs go to the reproductive tracts and the genital organs.
'•' ' / • '
i •;' Oc: - ' jr-" ' '> .
\ •ft^
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FIGURES
F ig , (1) Bra in of Danaus ch rvs ippus showing v a r i o u s
p a r t s : Pars i n t e r c e r e b r a i l s ( P i ) , Calyces
(Cy), S t a l k ( s ) . Corpora ped iuncu la t a (Cp).
F i g . (2) L a t e r a l s e c t i o n of t h e b r a i n .
2-6IJ
l-Bid
Fig, (3) Brain and the op t i c ganglion
connective (Cons),
Pig, (4) Optic ganglion with d i s t i n c t laminal
regions as seen under the l i g h t microscope,
j Brain (Br) , op t i c lobe (Opl) lamina (L),
medulla (M),
Fig-3
Fig.4
Fig, (5) Frontal sect ion of the thorac ic
ganglion with meso- and metathoracic
ganglion, Meso-thoracic ganglion (mes),
meta- thoracic ganglion (met), i n t e r -
ganglionic connective (Igc)
Fig. (6) Longitudinal sect ion of the f i r s t
thorac ic ganglion.
r 9
L
" " • " ^ ^ ^
;f m e s 'M
• • ' 1 . . « . • • , " ; — - •
k • ^ ' • ' ^ • • v ' ."*:
met ». /
J Fig.5
Flg-6
Pig. (7) Longitudinal section of the two
thoracic ganglia showing connection
with each other.
Pig. (8) Frontal view of metathoracic
ganglion (met).
Fig.7
Fig.8
Fig, (9) Longitudinal section of the thoracic
ganglion under higher magnification.
Fig, (10) Longitudinal section of first thoracic
ganglion showing neuromeres (Nm)
Fig.9
Fig 10
Pig. (11) Longitudinal sect ion of the raeso-
and raeta- thorac ic ganglion showing
ganglionic core .
Pig. (12) Frontal sect ion of meso- and meta-
thorac ic ganglia i n high magnification.
Fig.n
Fig.12
Pig. (13) Longitudinal section of the third
abdominal ganglion with neurilenuna
(Nlrn) and ganglionic core (gc)
Fig, (14) Longitudinal sections of third abdominal
ganglion in high magnification.
Fig-13
• : ^ -
FigU
Fig, (15) Diagram of the Central Nervous System showing
protocerebrum (Pr ) , deutocerebrum (dc),
t r i tocerebrum ( t r ) , thoracic ganglion (Tg)
abdominal ganglion (abd, gng. ) . Pars i n t e r -
ce rebra t i s ( P i ) , Optic nerve (Opt, ne r ) ,
antennal lobe (Ant, l ob ) , suboesophageal
ganglion (S .O.g , ) , Prothoracic ganglion
(Pro, gng.) 1st thorac ic ganglion (1 Tho,
gng), second thorac ic ganglion (2,Tho,gng),
th i rd thorac ic ganglion 3 , Tho gng)
th i rd ganglion (3rd Gng), fourth ganglion
(4th gng), f i f t h ganglion (5th Gng),
abdominal sp i racu la r nerve (abd. Spir , ner)
s ixth ganglion (6th Gang,), Genital chamber
(Gen. Cha). respec t ive ly .
Pa<S lA^.
roto. cer. to-cer.
3 rd Gng.
Ath Gng.
5th Gng.
obd. spir.ner.
Bth. Gng.
F'gis
%,
Fig. (16) Photomicrograph of the Central Nervous
systan showing Brain (Br,), thoracic
ganglion (Tg) and abdominal ganglion
(abd. gng).
Pig. (17) Abdominal ganglia with prominent
nerves.
Fig-ie
Fig.17
Fig. (18) Transverse section of abdominal
ganglia - Ganglionic core (gc).
Pig. (19) Photomicrograph of second abdominal
gangl ion ' I l n d abd, gng) .
Fig-18
Fig.19
Fig, (20) Thoracic gangl ion with ne rves . Thoracic
connec t ive (Tho, Con), P ro tho rac i c gangl ion
( P r o t . Gng), P r o t h o r a c i c connec t ive
( P r o t . Con), Mesothoracic ganglion (Meso.
Gng), Meta thorac ic gangl ion (Meta Gng.) L
L a t e r a l nerve (Lat , ner) v e n t r a l neirve Cord
(V.N.C. ) .
F ig , (21) Diagram showing t h e l a s t abdominal
gangl ion . Vent ra l nerve cord (V, N, c ) .
S ix th gangl ion (Vl th gng) nerve of
e x t e r n a l gangl ion (ne r , ex t . gng).
Tho.Con.
Prot.Gng.
Prot.Con.
Meso.Gong.
Meta.Gng.
FJ9.2O
V.N.C
Vlthgng.
ner. ext. gng.
Fi9-2I