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Pharmacological profile of octopamine and 5HT receptors on the lateral oviducts of the cockroach Periplaneta americana.

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Archives of Insect Biochemistry and Physiology 28:49-62 (1 995)
Pharmacological Profile of Octopamine and
5HT Receptors on the lateral Oviducts of the
Cockroach, Periplaneta americana
Shernaz X. B a m j i and Ian Orchard
Departmiit of Zoology, University of Tororito,Toronto, Ontario, Cariada
The effects of the amines SHT and octoparnine on the myogenic activity of
F'eriplaneta arnericana (L.) oviducts and the pharmacological profile of octopamine
and 5HT receptors on the lateral oviducts have been determined. Application
of 5HT to the oviducts resulted in a dose-dependent increase in basal tonus
and amplitude of contractions. Antagonist studies revealed that the 5HT receptor on the cockroach oviduct most resembles the mammalian 5HT2 receptor.
Application of octopamine resulted in a decrease in basal tonus and had a
biphasic effect on the amplitude of contractions, being stimulatory at low doses
and inhibitory at higher ones. The inhibitory effects of octopamine appear to
be mediated via CAMP and are blocked by antagonists which indicate that the
o 1995 Witey-Liss, Inc.
octopamine receptor is of the octopamine-2 type.
Key words: octopamine, SHT, oviducts, pharmacology, Periplaneta americana, visceral
The control of contractions of insect visceral muscles has been the subject of considerable interest for many years. The discovery of multiple
myotropic peptides and amines that affect insect visceral muscle (e.g.,
Holman et al., 1986; Orchard and Lange, 1985a,b; Robb et al., 1989) has
intensified the search for an understanding of the physiological regulation of muscle function.
The amines 5HT and octopamine have been implicated in the function of visceral muscle in a variety of insects, and evidence suggests that one or both of
these amines exert their effects on oviduct muscle, including Locusta rnigmtoria
Acknowledgments: This work was supported by the Natural Sciences and Engineering Research
Council of Canada and by insect Biotech Canada, a Federal Network of Centres of Excellence.
We are grateful to Agriculture Canada, London, ON, for provision of cockroaches.
Received September L O , 1993; accepted April 27, 1994.
Address reprint requests to Ian Orchard, Department of Zoology, University of Toronto, 25 Harbord
St., Toronto, ON, Canada, M5S 1 A l .
0 1995 Wiley-Liss, Inc.
Bamji and Orchard
oviducts (Orchard and Lange, 1985a,b), Tabanus sulcifrons oviducts (Cook
and Meola, 1978), and the oviducts of the cockroach, Periplanefa arnericana
(Stoya and Penzlin, 1988). Recently, the pharmacological properties of
octopamine receptors lying on the lateral oviducts of Locus t a migratovia
were examined (Orchard and Lange, 1986). Since the Periplaneta oviduct
responds to many peptides and amines in a similar manner to that of
Locusta migratoria, the cockroach oviduct is an ideal tissue upon which to
characterize and compare the pharmacological properties of both 5HT and
octopamine receptors with those found in other tissues.
5HT has been shown to be present in the nervous tissue of insects using
immunohistochemistry and quantitative electrochemical detection (see
Nassel, 1988), and although 5HT has been shown to be present in both
central and peripheral nervous systems as well as in neurohemal areas,
there is still relatively little known as to its physiological function in insects. Though extensive pharmacological work has been performed on
mammalian 5HT receptors, very little is known about insect receptor subtypes, and typically insect 5HT receptor subtypes have been classified with
respect to the criteria determined in mammalian systems. Antagonist studies have shown that in invertebrates the 5HT receptor subtype has most
often been described as being similar to that of the mammalian 5HT2 receptor subtype (Banner et al., 1987; Barrett and Orchard, 1990; Baines and
Downer, 1991; Barrett et al., 1993).
Conversely, the amine octopamine has been found in the nervous tissue of
every insect examined, and its role as a neurotransmitter, neurohormone, and
neuromodulator is well established (Evans, 1980; Orchard, 1982; Orchard and
Lange, 1987a,b).Although octopamine appears to be physiologically involved
in many insect systems, there are only a limited number of tissues which
have been shown to be innervated by identified octopaminergic neurons.
These include the lantern of firefly (Christensen et al., 1983), skeletal muscle
in the extensor-tibialis of locusts (Evans and O’Shea, 19781, and visceral
muscles in the oviducts of locusts (Orchard and Lange, 1985a,b).All of these
tissues receive information from dorsal unpaired median (DUM) neurons.
Many of the physiological functions of octopamine, including those in the
above three cases, appear to be mediated by a class of octopamine receptors
(octopamine-2 receptors) specifically linked to an adenylate cyclase (Nathanson, 1979,1985; Evans, 1984; Lange and Orchard, 1986).
The present paper examines the pharmacology of 5HT and octopamine receptors on cockroach oviduct and compares them to receptors in other tissues.
Adult female Periplaneta americana (L.)of varying age and reproductive state
were used throughout the study and were obtained from a long-standing
colony at Agriculture Canada (London, Ontario). The animals were reared
under crowded conditions at 25°C on a 12 hour light:12 hour dark photoperiodic regime and were fed rolled oats, dog chow, and Pablum baby food.
Amines and Cockroach Oviduct
Preparation of Tissue
Insects were dissected dorsally under cold physiological saline (150 mM
NaC1; 5 mM KC1; 5mM CaC1,; 2 mM MgC1,; 4 mM NaHCO,; 5 mM HEPES,
pH 7.2; 22 mM glucose). The abdominal sclerites, hindgut, and Malpighian
tubules were removed to expose the ovaries and oviducts. The preparation
for bioassays included the lateral oviducts and the common oviduct. For radioimmunoassay (RIA) only the lateral oviducts were used, whereas for immunohistochemistry, the sixth abdominal ganglia and the lateral oviducts
attached to the common oviduct were used.
For bioassays, the preparation was placed in a trough, containing 400 yl of
saline, moulded into the wax base of a dissecting dish. The posterior end of
the common oviduct was pinned to the base of the trough, and the anterior
end of the lateral oviducts was attached, using a piece of human hair, to a
force transducer. The preparation was mounted at an angle of approximately
45" to the horizontal by its attachment to the force transducer and was held
under a tension of approximately 50 mg. Myographs were recorded using a
transducer coupled with a pen recorder. A 200 y1 drug sample (at twice the
final concentration) was applied just under the surface of the saline bath while
200 yl of the saline was removed, maintaining the bath volume at 400 pl. The
response to the drug was recorded, and the preparation was washed several
times before the addition of the next drug.
The change in basal tonus induced by 5HT and octopamine was represented as a percentage of the maximum response seen with each respective
chemical. The data in each run was converted into a percentage of the maximum response in that run. The mean and standard error of each concentration was then calculated. Due to the fact that the maximum response was
observed at different concentrations in each run, no single concentration of
the dose-response curve necessarily represents 100% maximum response.
The change in amplitude of spontaneous contractions induced by 5HT and
octopamine was represented by the ratio of the amplitude of contractions
after the respective amine was applied, to the amplitude of contractions prior
to application. Hence, the value 1.0 indicates that no change in amplitude
was noted when the amine was added.
Radioimmunoassay for cAMP
The lateral oviducts from each cockroach were dissected out and placed
as a pair in saline containing 10 yl of 5mM 3-isobutyl-1-methylxanthene
(IBMX) and 10 pl of each of the appropriate pharmacological agents to a
final volume of 100 pl. The tissues were incubated for 10 min at room
temperature. Boiling 0.05 M sodium acetate buffer (500 yI) (pH 6.2) was
then added, and the mixture was boiled for a further 5 min. After a freezethaw, the tissues were sonicated and centrifuged for 15 min. Cyclic AMP
was determined in 50 yl aliquots of the supernatant using a commercially
available cAMP RIA kit (New England Nuclear, Lachine, Quebec). Results
were expressed as pmol of cAMP per sample, where the sample represented one lateral oviduct.
Bamji and Orchard
Immunohistochemistry and Electrochemical Detection
Oviducts and sixth abdominal ganglia were processed for immunohistochemistry using anti-serotonin antibodies as described previously (Lange
et al., 1988).
The presence of 5HT and dopamine in the oviducts was determined using
HPLC coupled with electrochemical detection as described previously (Orchard, 1990; Lange et al., 1988; Ali et al., 1993).
Effect of 5HT and Octopamine on Oviduct Contractions
Cockroach oviducts are myogenically active, as judged by the persistence
of phasic contractions when isolated from the central nervous system. It has
been previously shown (Stoya and Penzlin, 1988) that 5HT (serotonin) modulates phasic contractions and increases basal tonus. Stoya and Penzlin (1988)
also demonstrated that exposure of the Periplaneta arnericana oviduct to
octopamine results in a decrease in basal tonus and frequency and amplitude
of the spontaneous myogenic contractions. The effects of 5HT and octopamine
on the myogenic contractions of the cockroach oviduct were confirmed here
a5 a preliminary to defining the pharmacological profile of 5HT and
octopamine receptors located on the lateral oviducts of the cockroach.
5HT. Applications of 5HT resulted in a dose-dependent increase in basal
tonus and amplitude of phasic contractions of the myogenically active oviduct, although no significant changes in the frequency of myogenic contractions
were observed. The threshold for the change in amplitude lay between 10” M
and 5 x 10-*M,with a maximum effect (2.5-fold increase) at about 5 x
M to 5
x lo4 M (Fig. lA,B). The threshold for change in basal tonus lay between lo4 M
and lo-’ M, and the maximum change in the basal tonus was observed at 5 x
lo4 M (Fig. 1C). Interestingly, high doses of 5HT (5 x lo4 M and above) were
less effective than lower doses, possibly as a result of desensitization.
The actions of 5HT were inhibited by a variety of vertebrate serotonergic
antagonists. The amplitude of the basal contraction induced by
was compared to the amplitude induced by
M 5HT plus the respective
antagonist (Fig. 2). Cyproheptadine inhibited the actions of 5HT by an average of 58%, the largest inhibition seen of all the antagonists (Fig. 2A,C).
MDL72222 and mianserin inhibited the actions of 5HT by approximately 28%.
Propranolol was ineffective, and gramine and ketanserin (Fig. 2B,C) potentiated the effects of 5HT by 76% and 72%, respectively. The effects of these
antagonists alone on the spontaneous myogenic contractions of the oviduct
were examined, and no discernable change in the contractions was observed.
Octopamine. The biogenic amine octopamine had a biphasic dose-response curve for its effects on the amplitude of myogenic contractions (Fig. 3).
The amplitude typically increased at octopamine doses of 10-’0M to lo-’ M and
then decreased at concentrations of lo-’ M or greater. Examples of this are shown
in Figure 3A,C. The increase in amplitude of spontaneous contractions varied
between preparations, resulting in the rather large standard errors (Fig.3C).Thus,
we found difficulty in defining this response using antagonists. At concentra-
Amines and Cockroach Oviduct
lo-' lo-'
5HT Concentration (M)
log scale
lo-' m7
5HT Concentration (M)
log scale
Fig. 1. Response of the oviduct to 5HT.A: The response profile to increasing amounts of the
amine (applied at arrows). B: Dose-response curve for 5HT showing the change in amplitude of
the contractions as a function of concentration. The change in amplitude is the ratio of the
amplitude of the contractions in the presence of 5HT relative to the amplitude of the contractions in saline. C: Dose-response curve for 5HT showing the change in basal tonus expressed as
a percentage of the maximum obtained with 5HT on each individual preparation. Each point in
both 6 and C i s the mean of six experiments, and the vertical lines show the SE of the mean.
tions greater than
M, octopamine routinely resulted in a reversible relaxation of the basal tonus as revealed by a lowering of the baseline (Fig. 3A,B,D)
and an apparent decrease in the frequency of myogenic contractions. The vertebrate a-adrenergic receptor antagonist, phentolamine, completely blocked the
inhibitory effects of octopamine (Fig. 3B), as did mianserin.
Effects of 5HT and Octopamine on cAMP Content
Radioimmunoassays were used to quantify the cAMP levels of lateral oviducts incubated for 10 min in 5HT or octopamine.
Bamji and Orchard
i10-5M 5 H T
I O - ~~MH T
5 ~ 1 0 -ketanserin
Fig. 2. A,& Comparison of the response of the oviduct to lo-' M 5HT (traces on the left) with
the response of the oviduct to lo-' M 5HT in the presence of 5 x lo-' M cyproheptadine or
ketanserin (traces on the right). Each pair of figures was derived from the same preparation, and
the experiments were repeated on several different preparations. C: The change in amplitude of
oviduct contractions with lo-' M 5HT plus 5 x 10" M antagonist compared to the change in
the amplitude of contractions with 1 0-5 M 5HT alone. Each bar is a mean of five experiments,
and the vertical bars represent the SE of the mean.
Amines and Cockroach Oviduct
1 0 4 octopamine
10 M octopamine +
10 Mphentolamine
z1 1.2
5 0.2
5 -60
Octopamine Concentration (MI
log scale
Octopamine Concentration (MI
log scale
Fig. 3. Response of the oviduct to octopamine. A: The response profile to increasing concentrations
of the amine (arrows). B: The actions of lo-' M octopamine (left trace) is blocked by the presence of
lo-' M phentolamine (right trace). C: Dose-response curve for octopamine showing the change in
amplitude of the myogenic contractions as a function of concentration. The change in amplitude is
the ratio of the amplitude of contractions with octopamine to the amplitude in saline. D: Doseresponse curve for octopamine showing the change in basal tonus expressed as a percentage of the
maximum relaxation response obtained with octopamine for each individual preparation. Points in
both C and D represent the mean of six experiments, and the vertical lines show the SE of the mean.
Bamji and Orchard
Incubation of oviducts in 5 x lo4 M 5HT (a dose resulting in maximal
physiological effect) in the presence of IBMX revealed no change in the cAMP
content (not shown).
Incubation of oviducts with octopamine in the presence of 5 x lo4 M IBMX
resulted in a dose-dependent increase in cAMP content. A sigmoid relationship between the log of octopamine concentration vs. the increase in cAMP
content is evident in Figure 4. The threshold for cAMP elevation occurred
M octopamine with maximal elevation (twelvefold
between lo-’ M and 5 x
increase) occurring at about 5 x 10” M octopamine, thereby matching the
dose-response curve for inhibition of contraction by octopamine, rather than
the stimulation of contraction seen at lower doses.
The ability of a wide range of aminergic antagonists to inhibit octopamineinduced elevations of cAMP was examined to further characterize the receptor type. The addition of 5 x
M of the respective antagonists, in the
presence of IBMX, to the oviducts produced no decrease in cAMP levels, suggesting that endogenous octopamine was not stimulating the receptor (Fig.
4). Phentolamine inhibited the octopamine-induced increase in cAMP levels
by 87.3%, mianserin by 71.176, metaclopramide by 52.376, cyproheptadine by
38.5%, and gramine by only 20.7%. That is, the rank order of potency was
phentolamine > mianserin > metaclopramide > cyproheptadine > gramine
(see Fig. 4).
Determination of the Presence of 5HT in Lateral Oviducts
The presence or absence of 5HT in axons innervating the oviducts or in
muscles of the oviduct was determined using immunohistochemistry as well
as HPLC coupled with electrochemical detection.
Immunoreactive cell bodies for 5HT-like material were seen throughout
the sixth abdominal ganglia (not shown). However, neither the common oviduct nor the lateral oviducts contained 5HT-like immunoreactivity. To confirm the negative result obtained with immunohistochemistry, the presence
or absence of 5HT in the oviducts was determined using HPLC coupled with
electrochemical detection. The presence or absence of dopamine in the oviduct was also determined. No peaks were found to comigrate with either
5HT or dopamine (Fig. 5), even when the tissue was spiked with 5HT or
dopamine. This implies the absence of 5HT and dopamine in both the oviduct muscle and in terminals of axons innervating the oviducts.
Applications of 5HT to cockroach oviducts revealed a dose-dependent increase in basal tonus and amplitude of the myogenic contractions of the oviducts, with no significant change in the frequency of contractions. This effect
of 5HT on cockroach oviduct was also observed by Stoya and Penzlin (1988),
with the exception of the frequency of myogenic contractions, which they
felt increased with applications of 5HT but which in our study showed no
significant change in frequency. The oviduct musculature of the horsefly
Tabanus sulcifrons also exhibited an increase in muscle tonus and frequency
and amplitude of myogenic contractions (Cook and Meola, 1978), although,
oct. +metaclo.
oct. +cypro.
0 ,
CAMP dontent (prnol/@ample)
Bamji and Orchard
Fig. 5. A: Profile of dopamine and 5HT standards (1 prnol) using HPLC coupled with electrochemical detection. Dopamine is shown to run at 5.6 min and 5HT at 12.41 min. 6: Profile of
an extract of lateral oviducts from one cockroach. No peaks were found to co-migrate with
either 5HT or dopamine. Detection limits were approximately 50 fmol.
interestingly, neurally evoked contractions of locust oviducts were little affected by applications of 5HT (Lange and Orchard, 1984).
The effectiveness of various vertebrate serotonergic antagonists on the 5HTinduced change in amplitude, in order of increasing potency, was shown to
be cyproheptadine (58% inhibition) > MDL 72222 (28%inhibition) = mianserin
(28% inhibition) > propranolol (0% inhibition). Both gramine and ketanserin
potentiated the effects of 5HT, by 76% and 72%, respectively. This contrasts
Amines and Cockroach Oviduct
with the situation in the isolated locust foregut (Banner et al., 19871, where
ketanserin and mianserin were both effective competitive antagonists of the
5HT2-like receptor. Since the 5HT-mediated actions are most strongly influenced by 5HT2 receptor antagonists (i.e., cyproheptadine, mianserin, and
ketanserin), it would seem logical to conclude that the 5HT receptor of the
cockroach oviduct most resembles the 5HT2vertebrate receptor.
The mammalian 5HT1receptor is typically associated with increases in adenylate cyclase activity, whereas the mammalian 5HT2 sites are not. In the
present study 5HT failed to elevate cAMP levels, which is consistent with
the above conclusion. In invertebrates, the 5HT receptor subtype has most
often been described as being similar to that of the mammalian 5HT2 receptor subtype (Banner et al., 1987; Barrett and Orchard, 1990; Baines and Downer,
1991). However, in most of these and other reports, 5HT produced an increase in adenylate cyclase activity (Berridge, 1975; Banner et al., 1987; Barrett
and Orchard, 1990; Baines and Downer, 1991). For this reason, the cockroach
5HT receptor can be described as being similar to the mammalian 5HT2 on
the basis of antagonist studies but must be distinguished from other insect
5HT2 type receptors on the basis of mode of action. Though it may appear
curious that two 5HT antagonists, ketanserin and gramine, actually potentiate the 5HT-induced change in the amplitude of myogenic contractions, this
is not a unique observation. Lakoski and Aghajanian (1985) noted that in the
lateral geniculate nucleus of rats, ketanserin potentiated rather than attenuated the effect (in this case inhibitory) of 5HT.
There are differences between the properties of the cockroach 5HT receptor and those expected of a typical mammalian 5HT2 receptor. For example,
the antagonistic effect of MDL 72222, a 5HT3receptor antagonist, was equivalent to that of the 5HT2 receptor antagonist, mianserin. Consequently, it is
probably most accurate to classify this receptor as 5HT2-like,a complete comparison with a mammalian 5HT2receptor being difficult to achieve, and only
emphasizing that there are differences between vertebrate and invertebrate
Octopamine elicited a biphasic dose-response curve, increasing the amplitude of myogenic contractions at doses of lo-’’ M to lo4 M and then decreasing the amplitude at concentrations greater than
M. Octopamine was
observed to decrease the frequency of myo enic contraction and lower basal
tonus of the oviduct at concentrations of 10- M and greater. This observation
is consistent with results obtained by Stoya and Penzlin (1988), who also studied the effects of this amine on Periplaneta arnericana oviducts. In the locust
oviduct, octopamine inhibited both neurally evoked and myogenic contractions and lowered basal tonus in a dose-dependent manner (Orchard and
Lange, 1985a, 1986). The vertebrate a-adrenergic antagonist, phentolamine,
completely blocked the inhibitory effects of octopamine, with mianserin being less effective. Octopamine increased the levels of CAMP,and these effects
were found to be dose-dependent. Interestingly, the dose-response curve for
cAMP elevation matched the dose-response curve for inhibition of contractions induced by octopamine, suggesting that the stimulation of contraction
induced by low doses of octopamine is mediated via different receptors.
Octopamine also increased the cAMP content of lateral oviducts of Locustu in
Barnji and Orchard
a dose-dependent manner. Other insect systems in which octopamine’s actions are mediated by changes in cAMP include thoracic ganglia of Periplaneta
americuna (Nathanson and Greengard, 19731, the firefly lantern (Nathanson,
1979; Nathanson and Hunnicutt, 1979), brains of various insects (Bodnaryk,
1979; Harmer and Horn, 1977), locust flight muscle and oviducts (Worm, 1980;
Lange and Orchard, 1986), and locust and cockroach fat body (Orchard et al.,
1982; Gole et al., 1983).
The ability of a wide range of aminergic antagonists to inhibit octopamineinduced elevations of cAMP was examined to further characterize the receptor type. The rank order of potency was phentolamine (87.3% inhibition) >
mianserin (71.1% inhibition) > metaclopramide (52.3% inhibition) > cyproheptadine (38.5%)> gramine (20.7%inhibition). Characterization of the oviduct octopamine receptor using aminergic antagonists indicates its difference
from previously described octopamine receptors. The rank order of potency
of octopamine antagonists is comparable to but not identical to the rank order of antagonists on the Locusta oviduct which was found by Orchard and
Lange (1986) to be phentolamine > metaclopramide > mianserin > cyproheptadine > phenoxybenzamine > yohimbine > chlorpromazine.
The most detailed study of octopamine receptors has been made by Evans
(1981). Using a wide range of antagonists and agonists, Evans described the
presence of three distinct classes of octopamine receptors: octopamine-1 receptors which slow the rhythm of myogenic contraction and do not act via
cAMP and octopamine-2 receptors which modulate twitch tension, act via
CAMP,and can be further divided into type 2A receptors, lying presynaptically,
and 2B receptors, lying postsynaptically. The octopamine receptors mediating an
elevation in cAMP in cockroach oviducts are similar to the described octopamine2 receptors but do not conform to either 2A or 28 subtypes.
Orchard and Lange (1987a) determined by radioenzymatic assay that the
oviducts of Periplaneta americuna contain octopamine (31 pmol of octopamine
per mg protein). It would seem, therefore, that octopamine is a neurotransmitter /neuromodulator in the cockroach oviduct musculature, in a manner
similar to that shown for locust oviduct (Orchard and Lange, 1985a).Serotonin-like immunoreactivity was located in every ganglion in the CNS of the cockroach, although the largest number of immunoreactive cells occurred in the
terminal ganglion (Bishop and OShea, 1983).In the present study, cell bodies of
the sixth abdominal ganglia revealed serotonin-like immunoreactivity in accordance with the above results; however, there was no indication of serotonergic
innervation to the oviducts based upon immunoreactivity or HPLC coupled to
electrochemical detection. This would indicate that if 5HT plays a physiological
role in control of the lateral oviduct, then it is a neurohormone.
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cockroaches, octopamine, profil, periplaneta, lateral, 5ht, oviduct, receptors, american, pharmacological
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