In vitro and in vivo effects of myo-active peptides on larvae of the tomato moth Lacanobia oleracea and the cotton leaf worm Spodoptera littoralis (Lepidoptera; Noctuidae).
код для вставкиСкачать60 Matthews et al. Archives of Insect Biochemistry and Physiology 69:60–69 (2008) In Vitro and In Vivo Effects of Myo-Active Peptides on Larvae of the Tomato Moth Lacanobia oleracea and the Cotton Leaf Worm Spodoptera littoralis (Lepidoptera; Noctuidae) H. J. Matthews,* N. Audsley, and R. J. Weaver Neuropeptides from five different neuropeptide families [Manduca sexta allatostatin (Manse-AS), and Manse-AS deletion analogue5-15, M. sexta allatotropin (Manse-AT), leucomyosuppressin, perisulfakinin, and myoinhibitory peptide I (MIP I)] were assayed for their ability to affect the development and food consumption of penultimate and last larval instars of two lepidopteran species, L. oleracea and S. littoralis. Injections of Manse-AS deletion analogue5-15, Manse-AT, perisulfakinin, and MIP I had no observable effects on development, food consumption, or mortality compared to controls. Single injections of Manse-AS significantly reduced the weight gain and increased mortality of L. oleracea and S. littoralis larvae compared to controls. By contrast, feeding Manse-AS to L. oleracea had no such effects. These differences were probably due to the degradation of the peptide by digestive enzymes in the foregut of L. oleracea. In studies in vitro, perisulfakinin, and MIP I had no effect on the spontaneous foregut contractions of L. oleracea larvae. Leucomyosuppressin, however, had myoinhibitory effects on the foregut. Single injections of leucomyosuppressin significantly reduced the weight gain and food consumption of L. oleracea and S. littoralis larvae and increased mortality. These data suggest that the deleterious effects observed in vivo were due to the myoinhibition by Manse-AS and leucomyosuppressin of the normal peristaltic movements of the gut either by the intact peptide or by its cleavage products resulting from degradation in the haemolymph. Arch. Insect Biochem. Physiol. 69:60–69, 2008. © 2008 Wiley-Liss, Inc. KEYWORDS: Allatostatin; allatotropin; leucomyosuppressin; perisulfakinin; myoinhibitory peptide INTRODUCTION Lepidopteran pests cause damage largely through their feeding activities. Feeding and crop motility are controlled by the stomatogastric nervous system (Penzlin, 1985), and central to this system is the frontal ganglion, which, in Lepidoptera, controls the muscles of the foregut (Bushman and Nelson, 1990; Miles and Booker, 1994, 1998). Muscular activity of the foregut of larval Lepidoptera is regulated by the myoinhibitory allatostatins and myostimulatory allatotropin, which work together antagonistically (Duve et al., 1999, 2000). In larvae of the tomato moth Lacanobia oleracea, Manduca sexta allatostatin (Manse-AS) (pEVRFRQCYFNPISCF-OH), M. sexta allatotropin (Manse-AT) (GFKNVEMMTARGFNH2), and allatostatins of the Y/FXFGL-NH2 family are localized in the frontal ganglion, are detected in the recurrent nerve that innervates the muscles of the foregut, and have myoactivity on Central Science Laboratory, Sand Hutton, York, United Kingdom Contract grant sponsor: Pesticides Safety Directorate (DEFRA). *Correspondence to: H. J. Matthews, Central Science Laboratory, Sand Hutton, York, YO41 1LZ, UK. E-mail: j.matthews@csl.gov.uk Received 10 April 2007; Accepted 23 June 2008 © 2008 Wiley-Liss, Inc. DOI: 10.1002/arch.20265 Published online in Wiley InterScience (www.interscience.wiley.com) Archives of Insect Biochemistry and Physiology October 2008 Effects of Myo-Active Peptides on Lepidoptera the foregut (Duve et al., 2000; Audsley et al., 2005). In larvae of the cotton leaf worm, Spodoptera littoralis, a similar peptide profile has also been identified from the frontal ganglion (Audsley et al., 2005). Given the role of these neuropeptides in foregut motility, there is an interest in their use to suppress feeding of insect pests. Injection of ManseAS into L. oleracea larvae resulted in a reduction in larval growth and feeding, and increased mortality compared to control injected insects, most likely due the myoinhibitory effects of Manse-AS on the foregut. In contrast, injection of Manse-AT had no effect (Audsley et al., 2001). Manse-AS5-15 is a deletion peptide produced by cleavage of Manse-AS by haemolymph enzymes (Audsley et al., 2002b), and inhibits foregut contractions in the L. oleracea foregut in vitro at concentrations similar to those of the intact peptide (Audsley et al., 2001). As well as the allatostatins, other myoinhibitory peptides inhibit gut contractions in various insects. Six myoinhibitory peptides (termed MIP I–VI) were identified from the ventral nerve cord of M. sexta, and were shown to inhibit peristalsis of the anterior hindgut in adult M. sexta (Blackburn et al., 1995, 2001). The most abundant and the most potent of these peptides was MIP I (AWQDLNSAWamide; Blackburn et al., 2001). Leucomyosuppressin (pEDVDHVFLRF-NH2) is a FMRF-related peptide (FaRP) originally isolated from the cockroach Leucophaea maderae (Holman et al., 1986). It has since been identified in the cockroaches, Periplaneta americana and Blattella germanica (Predel et al., 2001; Aguilar et al., 2004), the honey bee Apis mellifera (Audsley and Weaver, 61 2006), and in species of Mantophasmatodea (Predel et al., 2005), but as yet, not in lepidopteran species. Leucomyosuppressin inhibits contractions of the foregut and hindgut in P. americana (Predel et al., 2001), L. maderaea (Cook and Wagner, 1991), and B. germanica (Aguilar et al., 2004), and inhibits spontaneous midgut contractions in the cockroach Diploptera punctata (Fusé and Orchard, 1998). When injected into adult B. germanica, it inhibited food intake in a dose-dependant manner (Aguilar et al., 2004). Perisulfakinin (EQFDDY(SO3H)GHMRF-NH2) is also a FaRP, isolated from the cockroach P. americana (Veenstra, 1989). Perisulfakinin displays sequence homology with vertebrate hormones of the gastrin/CCK family, which are satiety-inducing peptides (Lee et al., 1994). In the cockroach B. germanica, it induced foregut and hindgut contractions, and inhibited food intake when injected into the haemolymph (Maestro et al., 2001). Perisulfakinin inhibits food intake in nymphs of the locust, Schistocerca gregaria (Wei et al., 2000), although its mode of action was not determined. The aim of the present study was to compare the effects of various structurally different neuropeptides (Fig. 1) from different neuropeptide families on feeding and development in larvae of two species of lepidopteran pest insects, L. oleracea and S. littoralis. MATERIALS AND METHODS Insects Lacanobia oleracea and S. littoralis were reared as described by Corbitt et al. (1996) on an artifi- Fig. 1. Amino acid sequences and common names of peptides used in this study. Archives of Insect Biochemistry and Physiology October 2008 62 Matthews et al. cial diet and kept at 20°C, 65% relative humidity, in a 16-h light:8-h dark photocycle. Larvae were fed on a maize-based noctuid artificial diet (BioServ, Frenchtown, NJ). were also tested on S. littoralis only: S. littoralis physiological saline (Davenport and Wright, 1985), Dulbecco’s phosphate-buffered saline (Sigma), and M. sexta saline (Chamberlin, 1989). Peptides Injection Assays With Neuropeptides Manduca sexta allatostatin, deletion analogue Manse-AS5-15 and MIP I were custom synthesized at the Advanced Biotechnology Centre, Imperial College, London, UK. Manduca sexta allatotropin was purchased from Sigma-Aldrich (UK). Leucomyosuppressin and perisulfakinin were purchased from Bachem (UK). Prior to injection, Manse-AS and Manse-AS5-15 were dissolved in dimethyl sulphoxide (DMSO). Manse-AT, MIP I, perisulfakinin, and leucomyosuppressin were dissolved in water. Newly ecdysed fifth instar larvae (0–24 h old) of L. oleracea or S. littoralis were anaesthetized under CO2 and injected using a Hamilton syringe with either 3.8 µg (2 nmol) Manse-AS in 1 µl DMSO, 2.8 µg (2 nmol) Manse-AS5-15 in 1 µl DMSO, 1.5 µg (1 nmol) ManseAT in 1 µl water, 6.2 µg (5.7 nmol) MIP I in 1 µl water, 5.5 µg perisulfakinin (3.6 nmol) in 1 µl water or 18.9 µg (15 nmol) leucomyosuppressin in 3 µl water. Control insects were injected with the same volumes of either DMSO or water as appropriate. Following injection, larvae were weighed and placed individually in 250-ml plastic containers (Autobar Ltd, Hemel Hempstead, UK) lined with tissue paper and covered with a perforated lid. Larvae were fed artificial diet of known wet weight, and kept under rearing conditions. The weight of the larvae was subsequently monitored every two or three days until they either pupated or died. All data are means ± SE, n = 20–30. Food consumption was calculated by subtracting dry weight of remaining diet (dried to constant weight at 90°C) from converted dry weight of diet given to individual larvae. Dry weight of diet given to larvae was determined using a conversion factor calculated from the ratio of wet weight to dry weight of diet standards. Foregut Contraction Assay Sixth instar L. oleracea or S. littoralis larvae that had been starved overnight were anaesthetised with CO2 and dissected in a longitudinal well made in the wax base in a dissecting dish. The dorsal surface was opened by cutting from the level of the second proleg to the back of the head capsule, allowing exposure of the foregut and anterior midgut. The cuticular flaps were pinned down and the foregut washed several times with 200 µl of physiological saline of the following composition (mM): Na+ 154, K+ 2.7, Ca2+ 1.8, Cl– 160, hydroxyethylpiperazine ethanesulphonic acid (HEPES) 12, and glucose 22, pH 7.2 (Cook and Holman, 1978). The foregut preparation was then immersed in 200 µl saline. The number of peristaltic contractions were monitored over two 2-min periods to establish baseline contractions. Insects were discarded if there was no gut movement. Control saline was replaced with the same volume of solution containing either leucomyosuppressin at concentrations between 10–12 and 10–7 M, MIP I at concentrations of 10–7 and 10–6 M, or perisulfakinin at concentrations of 10–7 and 10– 6 M, and contractions monitored as above. Preliminary experiments had determined the most suitable concentration range. Ten different larvae were used for each concentration of peptide tested. The frequency of contractions in the presence of peptide was then compared to the frequency of control (baseline) levels. The following physiological salines Feeding Assays With Manse-AS Prior to feeding, Manse-AS was dissolved in DMSO. Newly ecdysed fifth instar larvae (0–24 h old) of L. oleracea that had been starved overnight were fed a small piece of noctuid artificial diet containing either 15 µg (7.9 nmol) Manse-AS in 1 µl DMSO or 1 µl DMSO only (controls). Larvae Archives of Insect Biochemistry and Physiology October 2008 Effects of Myo-Active Peptides on Lepidoptera 63 were left for 5 hours to consume the diet. Those that did not consume the whole piece were discarded from the experiment. Larvae were then weighed and subsequently monitored as for the injection assays. All data are means ± SE (n = 20). when tested in vitro, even with the addition of 10–8 M Manse-AT to stimulate contractions. Therefore, none of the peptides (leucomyosuyppressin, MIP I, or perisulfakinin) were tested in vitro on the foregut of S. littoralis. Statistical Analysis Effects of Neuropeptides on the Development and Survival of L. oleracea Larvae Dose-response results were analysed using probit analysis. Larval weights and diet weights were analysed using ANOVA followed by a multiple comparison test (Tukey’s) (significance level P < 0.05). Larval survival was analysed using the χ2 test (significance level P < 0.05). RESULTS Effects of Peptides on L. oleracea Foregut Peristalsis Leucomyosuppressin had an inhibitory effect on spontaneous contractile activity of the foregut of L. oleracea causing a complete inhibition at 10–7 M. The inhibitory effects were rapid in onset, and a dose-response curve for frequency of foregut contractions was produced between 10–12 M and 10–7 M leucomyosuppressin, giving an apparent EC50 of 2.2 × 10–11 M (Fig. 2). Neither MIP I or perisulfakinin influenced contractions of the L. oleracea foregut at doses up to 10–6 M. Effects of Peptides on S. littoralis Foregut Peristalsis None of the physiological salines tested on S. littoralis maintained foregut contractions in larvae In all experiments, control mortality was low, ranging from 0 to 16%. Following injection into fifth instar L. oleracea, Manse-AS had a significant effect on the subsequent growth and development of larvae. The mean weight of Manse-AS-injected larvae was significantly lower than that of controls (Fig. 3). In addition, the percentage mortality of treated larvae was significantly higher (45%) compared to controls. In contrast, feeding Manse-AS to L. oleracea had no effect on the subsequent weight of larvae or their mortality compared to controls. When leucomyosuppressin was injected into fifth instar L. oleracea, the weight of larvae post-injection was significantly reduced compared to controls (Fig. 4). Although not significantly different to controls, mortality of leucomyosuppressin-injected larvae increased to 32% at 14 days post-injection. Neither Manse-AS5-15, allatotropin, MIP I, or perisulfakinin had any significant effects on the development of larval L. oleracea at the doses tested. There were no significant differences between the weight of control and peptide-injected larvae. There were also no significant differences between survival of control and peptide-injected larvae in any of the above treatments. Fig. 2. Dose-response curve for the inhibition of peristalsis of the foregut of sixth instar L. oleracea larvae by leucomyosuppressin. Means ± SE, n = 10. Archives of Insect Biochemistry and Physiology October 2008 64 Matthews et al. Fig. 3. The effects of Manse-AS on L. oleracea and S. littoralis larval weight. Each histogram bar represents mean weight (mg) of larval L. oleracea or S. littoralis injected as day 1 fifth instar with either 3.8 µg Manse-AS in 1 µl DMSO or 1 µl DMSO (controls). Error bars represent standard errors. Effects of Neuropeptides on the Development and Survival of S. littoralis Larvae In all experiments, control mortality was low, ranging from 0 to 8%. The injection of Manse-AS into fifth instar S. littoralis larvae caused a significant reduction in weight gain compared to controls (Fig. 3). The survival of Manse-AS-injected larvae was also affected. Mortality was 32% in Manse-AS-injected larvae and this was found to be significantly higher than controls. Spodoptera littoralis larvae that had been injected with either allatotropin, MIP I, or perisulfakinin did not exhibit any significant effects on larval development compared to their respective controls. The weight of S. littoralis larvae from day 1 fifth instar through to pupation was not significantly affected by injection of the above peptides. Mortality of larvae post-injection was low in both allatotropin- and MIP 1–injected insects, ranging from 4 to 8%, and was not significantly different than controls. Mortality in perisulfakinin-injected insects was significantly higher than controls. Injection of leucomyosuppressin into fifth instar S. littoralis larvae caused a significant reduction in weight gain throughout their development to pupation compared to controls (Fig. 4). Mortality of larvae was also significantly higher for peptideinjected larvae (48%) compared to controls. Effects of Neuropeptides on Food Consumption of L. oleracea Lacanobia oleracea larvae that had been injected with either Manse-AS5-15, MIP I, or perisulfakinin Fig. 4. The effects of leucomyosuppressin on L. oleracea and S. littoralis larval weight. Each histogram bar represents mean weight (mg) of larval L. oleracea or S. littoralis injected as day 1 fifth instar with either 18.9 µg leucomyosuppressin in 3 µl water or 3 µl water (controls). Error bars represent standard errors. Archives of Insect Biochemistry and Physiology October 2008 Effects of Myo-Active Peptides on Lepidoptera did not show any significant differences in food consumption compared to controls. Feeding Manse-AS to fifth instar larvae had no significant effect on their subsequent food consumption through to pupation compared to controls. Injection of leucomyosuppressin into fifth instar L. oleracea caused a significant reduction in food consumption compared to water-injected controls. Food consumption was reduced in leucomyosuppressininjected larvae throughout their development to sixth instar (Fig. 5). At its maximum (day 9), each control larva had consumed an average of 275 mg food (dry weight) compared to 179 mg for leucomyosuppressin-injected larvae. Effects of Neuropeptides on Food Consumption of S. littoralis Spodoptera littoralis larvae that had been injected with either MIP I or perisulfakinin did not show any significant differences in food consumption post-injection compared to their controls. Injection of leucomyosuppressin, however, caused a significant reduction in the amount of food consumed by the larvae post-injection compared to water-injected controls. This reduction was apparent throughout their development to sixth instar (Fig. 5). The maximum amount of food was consumed at day 11, with an average of 216 mg (dry weight) per control larva compared to 171 mg per leucomyosuppressin-injected larva. 65 DISCUSSION Larvae of the two noctuid species L. oleracea and S. littoralis possess similar peptide profiles from their frontal ganglia. Three types of peptides were identified from frontal ganglion extracts; ManseAS, Manse-AT, and F/YXFGL-NH2 allatostatins, implicating them in the regulation of feeding activity (Audsley et al., 2005). This was confirmed in larval L. oleracea with Manse-AS, which inhibits foregut peristalsis in vitro and suppresses feeding activity causing mortality following injection into the haemolymph (Audsley et al., 2001). Results from the present study, which used a lower dose (3.8 µg Manse-AS per larva), showed similar but less pronounced effects, as might be expected given its dose-dependent effects (Audsley et al., 2001). Manse-AS was also biologically active following injection into larval S. littoralis at the same dose, causing a reduction in weight gain and increased mortality compared to controls. In vitro studies have confirmed the ability of high doses of ManseAS to inhibit foregut contractions completely in L. oleracea (Duve et al., 2000; Matthews et al., 2006; 2007) and it is probably this myoinhibition that is causing the effects on weight gain and mortality observed in this and other studies (Audsley et al., 2001). Although the in vitro activity of Manse-AS on the foregut of S. littoralis has not been shown, it is likely that Manse-AS is acting in vivo in a myoinhibitory manner in this species given the Fig. 5. The effects of leucomyosuppressin on L. oleracea and S. littoralis food consumption. Each histogram bar represents mean dry weight (mg) of cumulative food consumption by larval L. oleracea or S. littoralis injected as day 1 fifth instar with either 18.9 µg leucomyosuppressin in 3 µl water or 3 µl water (controls). Error bars represent standard errors. Archives of Insect Biochemistry and Physiology October 2008 66 Matthews et al. similarity of the results obtained. Injection of 1 nmol (ca. 1.9 µg) Manse-AS, however, twice daily into a different but related species, the fall armyworm S. frugiperda, had little effect on growth and development (Oeh et al., 2001). Although Manse-AT has myostimulatory effects on the foregut of L. oleracea (Duve et al., 2000; Audsley et al., 2005; Matthews et al., 2007), it did not affect development or food consumption when injected (this study; Audsley et al., 2001). Larval S. littoralis were also developmentally unaffected by this peptide, in contrast to the effects reported by Oeh et al. (2001), where twice daily injections of ManseAT into S. frugiperda larvae reduced weight gain and increased mortality in this closely related species. In addition, the duration of the penultimate and the last larval instars were prolonged in the survivors and injections of Manse-AT into adult female moths shortened the moths’ lifespan thereby reducing the total number of deposited eggs. The authors suggest that the effects of Manse-AT could have been caused by the intact peptide or by its cleavage products, since it was rapidly degraded by S. frugiperda larval haemolymph with a half-life ranging from 1 to 5 min (Oeh et al, 2001). Although Oeh et al. (2001) did not determine the mode of action of Manse-AT in vivo in S. frugiperda, its myotropic effects in vitro have been confirmed in other Noctuid species, for example Heliothis virescens (Oeh et al., 2003), H. armigera (Duve et al., 1999), and L. oleracea (Duve et al., 2000; Audsley et al., 2005). The contrasting results between different lepidopteran species are difficult to explain. They may largely be due to the different rates of degradation of the two peptides by haemolymph enzymes and/ or the relative activities of cleavage products resulting from this degradation. The degradation of Manse-AS by haemolymph from L. oleracea is rapid (half-life ca. 3.5 min), resulting in three major cleavage products (Manse-AS5-15, 6-15, 7-15; Audsley et al., 2002b). In vitro, only Manse-AS5-15 had any myoinhibitory activity on the foregut of L. oleracea larvae (Audsley et al., 2001; Matthews et al., 2006). It is, therefore, surprising that injection of the cleavage product Manse-AS5-15 had no effect on L. oleracea larvae in vivo. However, when injected, Manse-AS5-15 will be subject to degration by aminopeptidases in the haemolymph that sequentially cleave amino acids from the N-terminus (Audsley et al., 2002b). The removal of a single amino acid from Manse-AS5-15 would render this peptide inactive in vivo, because the truncated analogue ManseAS6-15 has no biological activity on the foregut of L. oleracea larvae (Matthews et al., 2006). This metabolism may occur before the truncated peptide can interact with the receptor, whereas the time taken for aminopeptidases to inactivate the intact peptide (Manse-AS1-15) by the sequential removal of five N-terminal residues, may enable an active truncated form of the peptide to interact with the receptors in the foregut. In contrast, feeding Manse-AS to L. oleracea larvae had no significant effects on growth, development, survival, or food consumption. This lack of measureable activity is most likely due to the enzymatic inactivation of the peptide in the gut and the ability of the peptide to penetrate the gut epithelium to reach its target receptors on the basolateral side of the gut. Audsley et al. (2002a) report that Manse-AS was rapidly degraded to Manse-AS4-15 and Manse-AS6-15 by trypsin-like enzymes associated with the foregut. Given that the truncated analogue Manse-AS5-15 has myoinhibitory activity, it would be reasonable to assume that the larger product, Manse-AS4-15 would also be active. However, this has not been tested in vitro. In the present study, leucomyosuppressin was the only other peptide tested, that had a significant effect on the growth and mortality of L. oleracea and S. littoralis larvae. This peptide has a very different structure to the allatostatins and it must be assumed that different receptors are responsible for its actions (Duve et al., 1995). Indeed, there is evidence from other species that this is the case (Cook et al., 1993; Vilaplana et al., 2004). Leucomyosuppressin had a powerful myoinhibitory effect on both the foregut and hindgut of adult B. germanica (Aguilar et al., 2004). The authors suggest that these myoinhibitory effects result in the accumulation of food in the foregut, which in turn inhibits food intake by the cockroaches because of the persistence of signals Archives of Insect Biochemistry and Physiology October 2008 Effects of Myo-Active Peptides on Lepidoptera from gut stretch receptors. Since leucomyosuppressin had a myoinhibitory effect on the L. oleracea foregut in vitro, this may explain the effects observed in vivo in L. oleracea and, by inference, in S. littoralis. Although perisulfakinin (like leucomyosuppressin) is a FaRP, it had no effect when tested in vitro on the foregut of L. oleracea or when injected into L. oleracea or S. littoralis. Perisulfakinin belongs to a family of insect neuropeptides known as the sulfakinins, which have myotropic and antifeedant effects (Maestro et al., 2001; Orchard et al., 2001). Mousley et al. (2004) found perisulfakinin to be inactive against the body wall muscle of the nematode Ascaris suum. They suggest that this could reflect the unusual structure of this peptide, possessing a sulphated tyrosyl residue unique to this subfamily of arthropod FaRPs. In the dipteran Calliphora vomitoria, perisulfakinin had no effect on crop contractions in vitro (Haselton et al., 2006). The authors propose that although it has myotropic and anti-feedant effects in insects other than P. americana, the amino acid sequence may be specific enough to prevent it from binding to blow fly alimentary tissue receptors on the crop, if present. These results might explain why this peptide had no effect on foregut peristalsis in L. oleracea in vitro or when injected into larval L. oleracea or S. littoralis. In addition, a lepidopteran sulfakinin has yet to be identified. All six myoinhibitory peptides isolated from the ventral nerve cord of M. sexta inhibit peristalsis of the anterior hindgut in vitro (Blackburn et al., 1995, 2001). Whilst MIP I was the most potent of these, it had no effect when assayed against the foregut of L. oleracea in vitro or when injected into L. oleracea or S. littoralis larvae. It is conceivable that receptors for MIP I are not present in the foregut, or that when injected this peptide is rapidly inactivated by haemolymph enzymes. ACKNOWLEDGMENTS The authors are very grateful to Hannah Bradish for the supply of insects used in this study. In addition, we acknowledge DEFRA for supplying a Archives of Insect Biochemistry and Physiology October 2008 67 license to obtain and keep S. littoralis (License no. PLH251C/5491(10/2006)). LITERATURE CITED Aguilar R, Maestro JL, Vilaplana L, Chiva C, Andreu D, Bell&eacaute;s X. 2004. Identification of leucomyosuppressin in the German cockroach, Blatella germanica, as an inhibitor of food intake. Regul Pept 119:105–112. Audsley N, Weaver RJ. 2006. 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