AN AUTONOMOUS POST GRADUATE TEACHING

DEPARTMENT OF ZOOLOGY AT MAHATMA JYOTIBA

PHULE EDUCATIONAL CAMPUS UNDER RASHTRASANT
TUKDOJI MAHARAJ NAGPUR UNIVERSITY

NAGPUR – 440033

SESSION 2023 – 2024

TOPIC – NEUROSECRETORY CELLS AND ORGAN OF

ODONATA

TEACHER IN CHARGE - SUBMITTED BY-

PROF. DR.S.C.MASRAM AISHWARYA D. ATILKAR
M.Sc ZOOLOGY (SEM-III)
DEPT. OF ZOOLOGY
RTM UNIVERSITY NAGPUR
 AN AUTONOMUS POST GRADUATE TEACHING
DEPARTMENT OF ZOOLOGY AT MAHATMA JYOTIBA
PHULE EDUCATIONAL CAMPUS UNDER RASTRASANT
TUKDOJI MAHARAJ NAGPUR UNIVERSITY

NAGPUR - 440033

SESSION 2023 - 2024

CERTIFICATE

THIS IS TO CERTIFY THAT I HAVE EXAMINED TEST OF

REVIEW ENTITLED ‘STUDY OF NEUROSECRETORY
CELLS AND ORGAN OF ODONATA

TEACHER INCHARGE
Prof. DR. S.C.MASRAM
SUBMITTED BY
AISHWARYA D. ATILKAR
M.Sc. (SEM-III)
DEPT.OF ZOOLOGY
Introduction

The nervous and endocrine systems work hand in hand in a process known as neuroendocrine
integration together, they play a crucial role in regulating various physiological functions,
ensuring the overall balance of the body, and responding swiftly to stress this collaborative
effort is essential for maintaining both everyday internal stability and a rapid reaction to
challenging situations (Hartenstein, 2006; Adamski et al. 2019).
A perusal of the ever-growing study of the neuroendocrine system of Insects has reveals that
little work has been done on Odonata in comparison to other orders the Present investigation
has, therefore, been undertaken to study the structure and Physiological activities of the
neuroendocrine system in the dragonfly Orthetrum Chrysis; this paper deals with the
histomorphology of the cephalic neuroendocrine system in the ultimate instar larva and the
adult (Highnam, 1969; Wigglesworth, 1970).

Since the initial documentation by day in 1940, extensive research has been conducted by
numerous scholars on the neurosecretory cells (NSC) located in the ventral ganglia of various
insects however, the majority of earlier investigations were limited to the NSC found in the
subesophageal ganglion more recently, there has been a shift towards more comprehensive
examinations, providing detailed insights into the cytomorphology, classification, and
distribution of NSC in not only the suboesophageal ganglion but also in the thoracic and
abdominal ganglia (Scharrer, 1941; Arvy, 1952).

Researchers have investigated neurosecretory cells (NSC) in Odonata, particularly focusing

on nymphs of certain dragonflies one notable study focus into the NSC not only in the
subesophageal ganglion but also extends the exploration to encompass the thoracic and
abdominal ganglia of nymphs belonging to the Aeschna grandis species this study
contributes to our understanding of the distribution and characteristics of NSC in different
parts of the nervous system among dragonfly nymphs (Charlet, 1969). Until now, there has
been a noticeable gap in research focusing on adult dragonflies study aims to shed light on
the intricacies of the structure and function of the neurosecretory system located in the ventral
ganglia of adult dragonflies, with a specific focus on the species Orthetrum chrysis the
neurosecretory mechanisms operating in the adult stage of this particular dragonfly species.
Histological studies and classifications of neurosecretory cells in dragonflies reveal there is a
notable absence of information regarding the nature of these cells specifically, there is no
account of whether they exhibit different phases of activity within a single cell or if they vary
from one another in their nervous system modules (NSM) , To address this gap, the current
study was initiated with the aim of determining the nature of neurosecretory cells in
Orthetrum Chrysis ( Schaller, 1968; Charlet, 1972).

The initial examination of the endocrine glands in Odonata, appears to have been inaccurate
in terms of the spatial arrangement of these organs. Baldus was the first to describe the nervi
corporum cardiacarum Interni (NCCI) and their connection with the "stomatogastric ganglia"
(corpora cardiaca) however, he did not recognize their functional significance. Furthermore,
Baldus provided an account of the protocephalic neurosecretory system in various Odonata
but couldn't find any indication of the nervi corporum cardi-Acarum externi (NCCII). He also
struggled to locate the nervi corporum allatarum (NCA) in nymphs it was reported that a
delicate pair of nerves, the NCCII, ran from the brain to the corpora cardiaca. Interestingly, it
is found that these nerves are present only in mature nymphs and adults. Additionally, Cazal
asserted the presence of nervi corporum allatarum (NCA) in mature nymphs this
information highlights the evolving understanding of the neuroendocrine system in Odonata
over different studies and time periods.

Brain

In the Odonata species, the neurosecretory cells within the brain showcase a cyclic pattern
of secretory activity during the developmental stage of the last instar larva notably , the
neurohormones released by the medial neurosecretory cells play a predominant role in
orchestrating the metamorphosis of the larva into a fully developed adult dragonfly this
information underscores the remarkable complexity of the neural structures in dragonflies
and sheds light on the crucial role played by neurosecretory cells and associated
neurohormones in the intricate process of metamorphosis from larva to adult (Schaller, 1959;
1970).
Larvae are treated with PME (Protocerebral medulla), the corpora cardiaca, which is a part
of the insect's brain, doesn't show any detectable neurosecretory material this is because the
corpora cardiaca functions as a neurohaemal organ, meaning it stores the neurosecretory
material produced by specific cells in the brain , In other words, instead of producing the
material itself, the corpora cardiaca acts like a storage facility for these important substances
that play a role in various physiological processes in the insect brain (Tembhare, 1976).
Larvae are treated with PME (Protocerebral medulla), it reduces the secretion activity of the
special cells within the corpora cardiaca in simpler terms , PME treatment decreases the
ability of specific cells in the insect's brain to release important substances stored in the
corpora cardiaca , this could have implications for various physiological functions in the
insect, as these substances play a role in regulating different processes within the body
(Varadaraj et al. 1993). The electrical activity of neurons responsible for detecting prey,
specifically those involved in recognizing downward movement on the opposite side (known
as descending contralateral movement detectors or DCMDs), is being studied in the brain. In
simpler terms, scientists are examining how certain brain cells respond to and process
information about the downward movement of potential prey understanding the
electrophysiological behavior of these neurons can provide insights into the mechanisms
underlying the insect's ability to detect and react to specific types of movement in its
environment (Gonka et al.1999).
Neurons in the dragonfly brain known as target-selective descending neurons (TSDNs). In
simpler terms, scientists are studying specific cells in the dragonfly's brain that play a role in
selecting and responding to particular targets these neurons are involved in the dragonfly's
ability to focus on and pursue specific objects or prey, Studying the behavior of target-selective
descending neurons helps us understand how dragonflies process information and navigate
their surroundings to capture their intended targets (Frye, 1995).

In the dragonfly species Orthetrum Chrysis, there are cells known as type B and C cells located
in the optic lobes of their brains these cells contain a substance that reacts positively to PF
staining it was observed specific cells in the dragonfly's optic lobes and these cells , labeled as
type B and C, show a positive reaction to a staining technique called PF staining this staining
indicates the presence of certain materials in these cells, Investigating these cells and the
substances they contain helps researchers better understand the structure and function of the
dragonfly's visual processing system ( Benedeczky. L et al. 1982 ). In the brain region called
the pars intercerebralis of the dragonfly Orthetrum Chrysis , there are three types of cells known
as A, B, and C cells these cells are distinct from each other and are responsible for producing
different types of neurosecretory material (NSM) to put it simply, in a specific part of the
dragonfly's brain , there are three different types of cells, labeled A, B, and C, and each of these
cells produces a unique chemical substance that plays a role in the insect's nervous system
investigating these distinct cell types helps scientists understand the diversity of functions
within the dragonfly's brain (Tembhare et al. 1975).

In dragonflies and damselflies (odonates), the corpora allata, which are glands involved in
hormonal regulation, have two different states these glands are actively producing hormones,
they are in a syncytial condition, meaning their cells are fused together without clear boundaries
on the other hand , when the corpora allata are not actively producing hormones, they are in a
cellular state with distinct cell walls separating individual cells these glands in these insects
have two forms: one where the cells are fused together during hormone production, and another
where the cells are separate when the gland is not actively engaged in hormone secretion this
distinction in the structure of the corpora allata is essential for understanding the hormonal
regulation processes in dragonflies and damselflies (Schaller et al. 1970; Tembhare et al.
1994).

Neurosecretory cells

In the ventral ganglia of insects, specifically in the lower part of their nervous system, there are
cells known as neurosecretory cells similar to the brain cells, these ventral ganglia cells come
in various types, and scientists classify them based on their preferences for different stains, as
well as their size and location in the ganglia this classification was established by (Raabe,
1971). The A, B, C1, and C2 cells in the ventral ganglia are specifically examined in terms of
their neurosecretory nature, meaning their ability to release substances that play a role in the
nervous system the fact that these cells produce chemically different nervous system modules
(NSM) provides strong support for categorizing them into distinct types of specific cells in the
lower part of the insect nervous system, and by studying their staining properties, size, and
location, they have found evidence supporting the idea that these cells function differently in
terms of the substances they produce for the nervous system (Thakare et al. 1975) .

The neurosecretory cells (NSC) found in the ventral ganglia of insects have similarities to those
located in the Pars intercerebralis region of the brain in Orthetrum chrysis (a type of dragonfly),
these cells responsible for releasing substances in the lower part of the insect's nervous system
are comparable to the cells found in a specific area of the brain in Orthetrum chrysis this
suggests that there are common features or functions between these two sets of cells, potentially
indicating shared roles in the regulation of the insect's physiological processes, Studying these
similarities helps scientists understand how different parts of the insect's nervous system work
together to control various functions in its body (Thakare et al. 1977) .

Cells labeled as A and B are considered genuine neurosecretory cells, meaning they truly
release substances that play a role in the nervous system on the other hand, cells labeled as C
are categorized as false neurosecretory cells, indicating that they don't actually release these
substances cells A and B, they actively contribute to the release of important substances in the
nervous system however, these cells labeled as C, they may resemble neurosecretory cells but
don't function in the same way; they don't release the substances associated with nervous
system regulation this distinction helps researchers understand the different types of cells and
their roles in the intricate functions of the nervous system (Highnam, 1961).

The B cells are identified as genuine neurosecretory cells because they contain certain chemical
groups, specifically SH (sulfhydryl) and SS (disulfide) groups and confirmed that B cells are
indeed neurosecretory cells by detecting the presence of specific chemical groups within them
these groups, SH and SS, are indicative of the cell's ability to release substances involved in
nervous system functions, this identification is crucial in understanding the role of B cells in
the intricate processes of the nervous system (Chalaye, 1967).

In the anterodorsal region of the medial part of the pars intercerebralis medialis, there are two
sets of neurosecretory cells located on either side of the root of the Medial ocellar nerve these
neurosecretory cells into three categories, labeled as A, B, and C, based on specific criteria this
classification was established by various scientists, including Nayar (1955), Johansson (1958),
Highnam (1961), and others, a particular area of the insect brain, there are cells that release
substances important for the nervous system. Scientists have observed these cells on either side
of a nerve root and have sorted them into three groups (A, B, and C) using certain criteria set
by researchers like Nayar, Johansson, Highnam, and others this classification helps in
understanding the diversity and functions of these neurosecretory cells in the insect brain
(Nayar, 1955; Johansson, 1958; Highnam, 1961; grillot, 1971).

Neurohormones –

Numerous neuropeptides undergo a process known as amidation at their C-terminus this

biochemical modification serves a crucial purpose – it effectively shields the free carboxyl
group present at the end of the peptide chain the rationale behind this protective measure lies
in preventing swift and indiscriminate degradation by carboxypeptidases when these
neuropeptides are circulating in the bloodstream to elaborate, amidation involves the
conversion of the terminal carboxyl group into an amide, a more stable chemical structure this
alteration is akin to placing a molecular guard on the vulnerable end of the neuropeptide,
shielding it from enzymatic attacks that could otherwise break it down too rapidly In the
dynamic environment of the bloodstream, where various enzymes are at play, this amidation
at the C-terminus acts as a strategic defense mechanism, ensuring the neuropeptides maintain
their structural integrity and functionality until they reach their intended targets or undergo
regulated degradation (Goldsworthy et al. 1989).

In the developmental stage of the dragonfly species Tramea virginia, specific cells known as
A-cells play a vital role these A-cells release a hormone called allatotrophic neurohormone,
which serves to activate the corpora-allata, and another hormone called prothoracicotropic
neurohormone, which stimulates the ventral glands this intricate hormonal signaling system
plays a crucial part in regulating various physiological processes during the larval stage of
Tramea virginia (Tembhare et al. 1994).
Neurohormones, which are signaling molecules in the nervous system, play a role in regulating
the Malpighian tubules, specialized structures involved in excretion in insects and other
arthropods when these neurohormones act on the tubules, they prompt a rapid release of fluid
from the hemolymph, the insect's circulatory fluid this heightened fluid movement leads to an
intensified diuretic response, where the insect expels a notably increased volume of urine this
physiological process is essential for maintaining the insect's internal balance of fluids and
waste products (Highnam, 1969; Cazal, 1947; Mordue, 1969).

The intrinsic cells responsible for regulating the sugar levels in the hemolymph of dragonfly
larvae do not produce neurohormones these intrinsic cells, which are integral to maintaining
the appropriate sugar concentration in the hemolymph (the circulatory fluid of the larva), do
not engage in the production of neurohormones these are signaling molecules that typically
play a role in the control and coordination of physiological processes in the absence of
neurohormone production by these intrinsic cells suggests that other mechanisms or factors
may be at play in the regulation of hemolymph sugar levels in dragonfly larvae physiology
remains a subject of interest and investigation in understanding the intricacies of metabolic
control in these developmental stages (Tembhare et al. 1991c).

Neurohormones, which are active substances made of proteins, are created and released by
regular neurons to elaborate, neurons, the fundamental cells of the nervous system, have the
capacity to produce and release neurohormones these neurohormones, being protein-based,
serve as signaling molecules that play important roles in transmitting messages within the
nervous system and often have widespread effects on various physiological processes the
synthesis and release of neurohormones by neurons are essential aspects of the intricate
communication network that enables the nervous system to regulate and coordinate activities
throughout the body (Bern, 1962; Scharrer, 1969; Wigglesworth, 1970).

Neurohormones, regardless of their origin, play a crucial role in maintaining the balance of
water and electrolytes in insects neurohormones are key players in the regulation of water and
electrolyte levels within the bodies of insects, Regardless of where these neurohormones come
from, whether it's the nervous system, certain glands, or other sources, their primary function
is to orchestrate processes that ensure a stable internal environment for the insect this includes
controlling the absorption and excretion of water and electrolytes, crucial factors for the insect's
overall physiological well-being the intricate involvement of neurohormones in these processes
underscores their significance in orchestrating the delicate balance of fluid and electrolyte
homeostasis in insect physiology (Novak, 1975).

Neuroendocrine system

Considerable information has been gathered regarding the MNC (Methyl Naphthyl Ketone)
groups in various other species of dragonflies, MNC groups in dragonflies beyond the specific
species initially mentioned this accumulated knowledge contributes to a broader understanding
of how MNC groups function across diverse dragonfly species, shedding light on their
ecological roles, behavioral patterns, or any unique features they may exhibit in different
contexts this wealth of information aids researchers in comprehensively studying and
comparing the roles and behaviors of MNC groups across the broader spectrum of dragonfly
species (Hanstrom, 1940).
The natural peptide antigens found in the stained neurons could be a part of either the pancreatic
polypeptide family or the FMR Famide family to delve deeper, when examining stained
neurons, scientists have identified endogenous peptide antigens these particular peptides, or
protein fragments, might be associated with two distinct families: the pancreatic polypeptide
family or the FMR Famide family these families are groups of related peptides with specific
structures and functions, Investigating the presence of these peptide antigens in neurons
provides valuable information about the potential involvement of these peptide families in
neural processes, offering insights into the complex molecular dynamics within the nervous
system (White et al.1986).

The regulation of ionic balance through hormones seems primarily associated with the
neurohormones produced by A cells, However in certain insects, the cells of the Pars
intercerebralis in the brain also play a role the intricate system of hormonal regulation that
maintains the balance of ions within an organism appears to have a primary player in the form
of neurohormones from A cells these neurohormones likely orchestrate the physiological
processes involved in ionic balance notably, in specific insects, an additional regulatory
component comes into play – the cells of the Pars intercerebralis located in the brain these cells
likely contribute to the overall control of ionic balance, showcasing the complexity and
variation in hormonal regulatory mechanisms across different insect species of brain (Berridge,
1966; Vietinghoff, 1966; Wall, 1967; Mills, 1967; Pilcher, 1970).

In B cell Geminata, A-cells exhibit active synthesis and secretion of neurohormones but in
PME treated larvae, the Cells gradually stop their secretory activity technics and radio
immunoassays of extracts from the brain, several gastro-entero-pancreatic hormones and
neuropeptides have previously been detected in the insect nervous system, e.g. :
cholecystokinin/gastrin (Dockray. J. G, 1981; Duve. H, 1982; El-Salhy. M, 1980).

Despite the absence of fully developed olfactory glomeruli, the dragonfly species L. depressa
demonstrates the ability to sense olfactory stimuli, as evidenced by electrophysiological
recordings L. depressa lacks fully mature olfactory glomeruli, which are specialized structures
in the brain associated with the sense of smell, the dragonfly displays proficiency in detecting
olfactory stimuli this capability has been observed and confirmed through the use of
electrophysiological recordings, a method that involves measuring electrical activity in
response to sensory stimuli the findings suggest that L. depressa employs alternative or less
conventional mechanisms to perceive and respond to olfactory cues, highlighting the
adaptability and complexity of sensory processes in this particular dragonfly species (Rebora
et al. 2012).

Distinctive features known as olfactory glomeruli are characteristic elements found in the
primary olfactory centers of both vertebrates and advanced invertebrates the olfactory
glomeruli are specialized structures found in the brains of vertebrates and certain higher
invertebrates, serving as crucial components of their primary olfactory processing centers these
structures play a key role in the initial stages of olfaction, where sensory information related to
smell is received, integrated, and processed the presence of olfactory glomeruli associated with
the sense of smell in both vertebrate and advanced invertebrate species (Hildebrand et al. 1997;
Joseph, 2020).
The internal head anatomy of Epiophlebia in its larval stage is severely incomplete and lacking
in detailed information while examining the internal structures of the head in the larval phase
of Epiophlebia, researchers have encountered significant challenges due to the fragmentary
nature of the available information the limited details hinder a comprehensive understanding
of the larval internal head anatomy of this particular species this gap in knowledge underscores
the need for further research and exploration to fill in the missing pieces and gain a more
thorough insight into the intricacies of Epiophlebia's larval head anatomy (Asahina, 1954;
Tramu. G et al.1983). Lead ions being effective in this connection could only be present In
very low concentrations in the hemolymph fluid, which normally is separated from The brain
tissue by a neural lamellar sheath as an effective barrier against uncontrolled Ion influx
(Treheme, 1972).

Conclusion

The adult dragonfly, Orthetrum chrysis, exhibits a classification of neurosecretory cells in its
ventral ganglia, denoted as A, B, C1, and C2 cells. These cells are identified based on three
criteria: the presence of stainable granules, the occurrence of neurosecretory material in their
axons and terminals, and cyclic activity that involves changes in the quantity of
neurosecretory material corresponding to physiological activities. A neuroanatomical study
has unveiled the existence of spherical knots in the aglomerular antennal lobe, suggesting a
potential role in the perception of odors. Neurons believed to regulate interception in
dragonflies are accessible through tracts in the third neuropil of the optic lobe. The
subesophageal ganglion (SEG), located ventrally just below the brain and esophagus, exhibits
the presence of immunoreactive deposits within the insect neurohaemal organ. The term
"cephalic endocrine glands" encompasses various structures, including neurosecretory cells
of the pars intercerebralis and their axons, corpora cardiaca, corpora allata, and ventral
glands.

The physiology of neurons accessible through tracts in the third neuropil of the optic lobe,
and the neurosecretory axons, likely corresponds to those immunohistochemically detected.
Interestingly, the removal of the pars intercerebralis region of the brain and ligature at the
neck region do not provide any indication of involvement of hormones from the brain and
other endocrine organs of the cephalic region. Flight muscles, which oscillate under direct
flight control, exhibit one-to-one matches between neuronal stimulus. Notably, in both adult
and nymphal dragonflies, the same brain regions seem capable of processing highly diverging
information from different sensory structures. This intricate network of neurosecretory cells,
anatomical structures, and physiological processes underscores the complexity and
adaptability of the dragonfly's nervous and endocrine systems.
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