沙柳木蠹蛾性行为及其性信息素滴度的动态节律

沙柳木蠹蛾Holcocerus arenicola是为害沙柳Salix psammophila的主要钻蛀性害虫之一。在进入暗期后的6 h内,对其求偶行为和交配行为进行观察和记录,调查该虫求偶和交配的活动规律。通过触角电位技术（EAG）和毛细管气相色谱（GC）对性腺体内信息素的滴度进行了分析,以揭示性信息素产生的昼夜节律和雌虫日龄对性信息素合成的影响。结果表明：该虫在暗期0.5~1 h内即开始有求偶行为,最大求偶高峰出现在羽化后的第2晚。交配行为主要发生在暗期的1~2 h内。在各日龄的成虫中,1日龄成虫的交配百分率最高。交配平均时间为24.16±2.64 min,随着日龄的增加,交配时间前移。在实验中,未观察到5~6日龄成虫的交配行为。沙柳木蠹蛾性信息素的体内合成早于求偶行为1~2 h,并在暗期的前2 h内达到峰值。性信息素的滴度随着雌虫日龄的增减而减少,最高值为当日羽化雌虫的腺体提取物。林间诱蛾实验中,处女雌蛾和性腺体提取物对雄蛾均有较好的诱捕效果。本研究表明,在沙柳木蠹蛾的性信息素滴度和性活动之间具有同步关系,同时为进一步利用长距离性信息素防治该虫提供了参考。     

竹笋基夜蛾的求偶及交配行为

在光周期14 L:10 D、温度(25±1)℃、相对湿度(60±10)％的室内条件下,研究了竹笋基夜蛾的求偶、交配行为,以及雄蛾对雌蛾腺体提取物的EAG反应.结果表明:竹笋基夜蛾求偶及交配行为仅发生在暗期.雌蛾羽化当天便可求偶,第2日雌蛾求偶率达最高;雌蛾进入暗期0～4h后开始求偶,5～7h求偶率达最高,暗期结束前1～2.5h终止;求偶的起始时间随着日龄的增加逐渐提前,1～4日龄雌蛾求偶的次数及持续时间随着日龄的增加而上升,5日龄则下降;雌蛾求偶高峰及停止求偶的时间随日龄的增加逐渐提前.成虫羽化当日便可交配,第2日达到交配高峰,5日后停止交配;雌蛾进入暗期5.5～7.0 h达到交配高峰,交配高峰出现的时间随日龄的增加而提前;随日龄的增加,交配起始时间逐渐提前,而交配持续时间逐渐缩短;增大雌雄比,竹笋基夜蛾交配起始时间提前,交配持续时间缩短,交配率显著提高.竹笋基夜蛾雄虫对2日龄处于求偶期雌虫的性腺体粗提物有显著的EAG反应.     

六星黑点豹蠹蛾求偶行为与性信息素产生和释放的时辰节律

六星黑点豹蠹蛾Zeuzera leuconotum Butler(Lepidoptera:Cossidae)是一种重要的园林害虫。为研究利用其性信息素防控六星黑点豹蠹蛾,在光周期L∶D=14∶10、温度19—32℃、相对湿度75%—85%条件下,对雌蛾求偶行为进行了观察;采用腺体提取法和瓶内收集法分别制备了不同日龄在同一时辰、同日龄在不同时辰的雌蛾性信息素粗提物,然后测定雄蛾触角对各种性信息素提取物的触角电位反应;以雄蛾对腺体提取物的EAG值表示性信息素的产生量,对瓶内收集物的EAG值表示释放量,检测性信息素产生与释放的时辰节律。结果表明:雌蛾求偶均发生在暗期;雌蛾求偶率与日龄有关,其中2日龄雌蛾求偶率最高,平均为86.7%;不同日龄雌蛾求偶高峰期均在进入暗期4—6h期间;随着虫龄的增加,求偶高峰期前移。雌蛾羽化当晚体内即可产生性信息素,并且向体外释放,但量较少;2日龄雌蛾产生和释放的性信息素量最大;雌蛾腺体中性信息素含量从进入暗期后逐渐增加,在暗期5h含量最高,随后逐渐减少;雌蛾释放性信息素从暗期4h开始,至暗期6h释放量最大。通过林间处女雌蛾诱捕试验证明了六星黑点豹蠹蛾的求偶行为与性信息素产生和释放在时辰节律上具有一致性。     

褛裳夜蛾的交配行为及雄蛾对性腺提取物的反应节律

详细观察了褛裳夜蛾成虫的交配行为,利用风洞、触角电位技术研究了雄蛾对性腺提取物的反应节律,通过林间诱蛾试验进行了验证,旨在为褛裳夜蛾性信息素的精确提取及性信息素组份分离、鉴定提供依据。研究结果表明:成虫的求偶、交配行为均发生在暗期,有一定的节律性:雌蛾在羽化3d以后开始求偶,1周左右表现最为强烈,3-4日龄雌蛾在暗期4-6h开始求偶,最大求偶率在暗期6-8h;5-9日龄在进入暗期就开始求偶,最大求偶率出现在暗期4-6h。风洞试验表明,3-7日龄的雄蛾对性腺提取物均有明显的性行为反应,5日龄雄蛾在暗期4-6h对性腺提取物的反应最为强烈。3-6日龄的成虫在暗期4-6h开始交配,而7-8日龄的成虫于暗期2-4h开始交配,6日龄的成虫交配率最高,交配高峰在暗期4-8h。雄蛾对性腺提取物的触角电位反应也有一定的节律性:雄蛾对4日龄雌蛾性腺提取物开始有电生理反应,对6-7日龄暗期5h性腺提取物反应最为强烈。林间诱蛾试验测定了性腺提取物的引诱活性,7日龄雌蛾性腺提取物林间诱蛾量最高,引诱高峰在暗期4-6h,该结果也验证了褛裳夜蛾雄蛾对性腺提取物的反应节律。     

竹织叶野螟成虫求偶行为及其雌蛾性信息素分泌腺超微结构

【目的】蛾类昆虫性信息素的合成和释放与求偶行为的发生是一致的,其合成和释放的器官是性信息素腺体。为深入了解竹织叶野螟性信息素的分泌生理,开展了竹织叶野螟求偶行为及其性信息素腺体超微结构的研究。【方法】在光周期14L:10D、温度26±2℃、相对湿度80%±10％的室内条件下,观察研究了竹织叶野螟的求偶行为;依据求偶规律研究结果,选取最活跃日龄雌蛾,在暗期求偶高峰时间段,充分挤压其腹部末端,然后于第8节处横向切下,将切下的腹末标本处理后,借助显微镜和扫描电镜观察性信息素腺体的表面特征及超微结构。【结果】竹织叶野螟雌雄蛾求偶均具一定的程序性,且求偶行为只发生在暗期,暗期前5 h内雌蛾求偶率较低,6 h后求偶率明显升高,并在暗期7-8 h达到求偶高峰;求偶率与雌蛾日龄有密切关系,3日龄雌蛾求偶率最高,持续时间也最长。竹织叶野螟性信息素分泌腺位于腹部第8-9节节间膜上,是一完整的环状结构,显微镜下观察其分泌腺为一乳白色囊状体,扫描电镜下其腹面囊状体迂回褶皱多,大体分为3个褶皱区,除第1褶皱区外,其余褶皱区表面密布乳突、凹陷沟和刺状物,且刺状物顶端有孔;背面囊状体皱褶少,其表面形态与第2和第3褶皱区相似。【结论】研究结果有助于了解竹织叶野螟性信息素合成和释放的时辰节律,也为该虫性信息素的准确提取和鉴定、性信息素的生物合成及利用提供了科学依据。     

光强度对棉铃虫交配行为的影响

【目的】为阐明光强度对棉铃虫Helicoverpa armigera (Hübner)交配行为的影响。【方法】本实验设置4个光照强度(0,0.5,5.0和50.0 lx),观察记录不同光照强度下棉铃虫雌蛾的求偶行为;分别通过单个腺体性信息素提取法和解剖雌虫受精囊的方式,检测不同光照强度下棉铃虫的雌蛾性信息素滴度和交配率。【结果】在强光(50.0 lx)下,棉铃虫雌蛾求偶起始时间最长,求偶持续时间最短,求偶次数最少,雌蛾性信息素滴度始终处于较低水平,交配率也属于最低。在微光(0.5 lx)下,雌蛾求偶起始时间最短,求偶持续时间最长,求偶次数最多;雌蛾性信息素始终处于较低水平,但暗期后段求偶率高达40%。【结论】强光(50.0 lx)可以抑制棉铃虫雌蛾的求偶行为、性信息素的合成及交配;微光(0.5 lx)可以促进棉铃虫雌蛾的求偶行为;相对黑暗环境(0 lx),微光(0.5 lx)还可以促进棉铃虫快速(1 h)完成交配,微光(0.5 lx)对棉铃虫的交配行为具有重要意义;求偶和性信息素的合成没有必然联系。本研究可为探讨光对夜蛾交配行为的影响提供一定的理论基础,也可为利用物理、化学通讯信息调控夜蛾行为提供一定的参考。     

笋秀夜蛾的求偶及交配行为

笋秀夜蛾(Apamea apameoides)是我国南方地区最为重要的笋期害虫,了解其求偶和交配行为是性信息素分离鉴定和应用的关键之一。在室内条件下,对笋秀夜蛾的性比、寿命和求偶、交配行为进行了研究。结果表明:笋秀夜蛾雌雄性比为1∶0.76;雌蛾和雄蛾的平均寿命因交配缩短;笋秀夜蛾的求偶和交配行为仅发生在暗期,日龄对其求偶和交配行为影响显著。雌蛾羽化当天便可求偶,第2日雌蛾求偶率达最高,求偶的起始时间随着日龄的增加逐渐提前。成虫羽化当日便可交配,第3日达到交配高峰;随日龄的增加,开始交配的时间逐渐提前,日龄对成虫交配高峰出现的时间影响不显著。     

棉大卷叶螟求偶行为节律和性信息素鉴定

[目的]观察棉大卷叶螟Sylepta derogata雌蛾的求偶行为及其动态节律,并鉴定该虫的性信息素组分,为开发该虫的性诱剂并应用于种群测报和绿色防控提供依据.[方法]将棉大卷叶螟处女雌蛾单头放入塑料杯中,从暗期开始前0.5 h到暗期结束后0.5 h,每隔0.5 h观察并记录1次雌蛾的求偶情况;在棉大卷叶螟雌蛾求偶高...     

豆野螟成虫行为学特征及性信息素产生与释放节律

豆野螟Maruca vitrata (Fabricius)是一种严重的泛热带豆类蔬菜害虫。本文在（29±1）℃、相对湿度75%～80%、光周期14L∶10D条件下研究了豆野螟成虫的羽化、交尾行为以及雌蛾性信息素的释放节律。结果表明：其羽化行为全天可见,在雌蛾中,86%于暗期羽化; 在雄蛾中,73%于暗期羽化。雌雄蛾羽化行为在暗期第4、5和8 h差异达到显著 (t>4; P<0.05)。交尾活动发生在暗期19：00到5：00之间,交尾持续时间最短约为20 min,最长约为90 min,3日龄进入暗期第5 h具有最高的交尾率。1、6和7日龄成虫具有单个交尾高峰,2到5日龄成虫具有两个交尾高峰。同一日龄成虫交尾在暗期前半段平均花费的时间要明显高于在后半段花费的时间。低龄和高龄的成虫用于交尾的时间明显高于中龄的性成熟成虫。成虫的开始交尾时间随着日龄的增加逐渐前移。雄蛾对进入暗期后第5 h和第9 h处女雌蛾的性腺提取物和空气收集性信息素的触角电位反应最强,对 3日龄处女雌蛾的性腺提取物和空气收集性信息素的触角电位反应最强。处女雌蛾田间诱蛾试验表明：23：00-01：00为诱蛾高峰期,3日龄处女雌蛾的诱蛾效果最好。该蛾的羽化、交尾及性信息素产生与释放均存在节律上的一致性。雌蛾的性信息素释放的时间较长,见于整个暗期,然而交尾行为发生时间较短,主要发生于两个交尾高峰之间。     

烟夜蛾雄蛾性附腺因子对雌蛾性信 息素合成的抑制作用

烟夜蛾Helicoverpa assulta处女蛾在交配后1 h,其性信息素滴度即显著降低,72 h内未见恢复。生测结果表明,烟夜蛾性信息素合成抑制因子主要来源于雄蛾性附腺。不同日龄雄蛾性附腺提取物的抑制活性无显著差异。光暗期对其活性具显著影响,暗期中雄蛾的性附腺物质对雌蛾性信息素合成具有较强抑制作用,而光期中雄蛾的性附腺物质不具抑制活性。在暗期的不同时间处理,对处女蛾性信息素合成的抑制作用无显著差异。雄蛾性附腺提取物对雌蛾性信息素合成的抑制作用与注射剂量有明显的相关性,0.2 ME（雄蛾当量）是产生显著抑制作用的最小剂量。对交配雌蛾注射性信息素生物合成激活神经肽（PBAN）提取物后,其性信息素合成又可恢复,这说明雌蛾交配后,性信息素滴度降低的原因是由于缺少了PBAN的调控。     

DIEL RHYTHMS OF CALLING BEHAVIOR AND SEX PHEROMONE PRODUCTION OF BEET ARMYWORM,SPODOPTERA EXIGUA (LEPIDOPTERA:NOCTUIDAE)*

Abstract Beet armyworm (BAW), Spodoptera exigua, is becoming one of more and more serious pests in China in recent years. As a part of research program of sex pheromone and its application of BAW in China, the hourly and daily variation of calling behavior and pheromone production of BAW females were investigated. Both calling behavior and titers of 4 sex pheromone components showed distinct diel rhythms, and the two peak periods were synchronous. In comparison, the calling activity lasted shorter period of time with a longer peak time, whereas the production of the sex pheromone lasted throughout the whole scotophase and part of the photophase with a very short peak time. The calling behavior began at the middle scotophase, reached the maximum at the middle‐later scotophase, and continued the maximal calling activity until the end of the scotophase. When the light was on, the calling percentage reduced sharply, and all females stopped calling 1 hour later. The variation patterns of the 4 pheromone components in the glands of the 3 day old moths were similar from one to another. From 0.5 h before to 4.5 h into scotophase, the titers increased slightly, but at 6.5 h they showed a significant increase up to the peak values of the scotophase. Soon at 8.5 h into the scotophase, they decreased significantly and thereafter gradually to undetectable level at 4.5 h into the next photophase. The daily change experiment showed that BAW females began calling at 0 scotophase, became fully active in calling at 3rd scotophase, and maintained the calling activity to 7th scotophase. There was no significant difference in pheromone titers among different day‐old moths.     

大螟成虫性信息素生物合成和释放及求偶和交配行为的昼夜节律

【目的】探索大螟Sesamia inferens性信息素顺11-十六碳烯乙酸酯(Z11-16∶Ac)和顺11-十六碳烯醇(Z11-16∶OH)的合成和释放及求偶和交配行为的昼夜节律,及其与田间性信息素诱捕的关系。【方法】通过溶剂浸提和固相微萃取(solid phase microextraction, SPME)分析大螟雌蛾性信息素Z11-16∶Ac和Z11--16∶OH的滴度,结合行为观测和多地田间实时性信息素诱捕数据,调查大螟性信息素的生物合成、释放及求偶和交配行为的昼夜节律。【结果】大螟雌蛾腺体内性信息素Z11-16∶Ac和Z11-16∶OH含量可检测到的时间始于暗期前1 h,暗期后4 h快速增加,暗期8 h为第1次高峰,但光期1h又一次高峰,光期5 h还可以被显著检测到。分泌至腺体外的性信息素化合物可检测到的时间始于暗期后6 h,高峰期在暗期后10 h,光期后1 h性信息素Z11-16∶Ac滴度达到96.9±20.9 ng/雌。采用溶剂浸提法获得的Z11-16∶Ac和Z11-16∶OH的比例在暗期平均为2.8±1.9,在光期平均为2.5±0.9,统计上二者没有显著差异,而SPME法获得的Z11-16∶Ac和Z11-16∶OH的比例在暗期平均为8.5±1.2,在光期平均为5.7±0.6,统计上二者差异显著。产卵器伸出时间发生在暗期6-8 h,产卵器伸出持续时间平均为80.8±4.4 min。大螟的交配发生在暗期4-10 h,交配持续时间平均为83.4±5.0 min。广东、四川、浙江、江苏四省性诱自动计数的田间每日每小时实时计数数据显示,越冬代诱蛾比较集中,之后的世代则比较分散,田间雄蛾的性诱昼夜节律受地理环境、季节和世代等因子的影响。【结论】本研究发现大螟交配和性信息素释放的昼夜节律在时间上不一致,交配时间在暗期较早时段。雌蛾性信息素有效的释放时间范围比雄蛾对性信息素反应的要小。产卵器伸展与雌蛾性信息素化合物的释放速率加快和扩散 范围有关。     

Calling Behavior and Rhythms of Sex Pheromone Production in the Black Cutworm Moth in China

The hourly and daily variation of calling behavior and sex pheromone production by the female black cutworm moth, Agrotis ypsilon (Rottemberg), were investigated. Both calling behavior and titers of five sex pheromone components showed distinct diel rhythms, and peak periods of calling and pheromone titer were synchronous. Females began to call on the first night after eclosion and called mainly during the second half of the scotophase. The percentage of females calling increased in 1 to 3-d-old females and decreased in 3 to 7-d-old females. Females advanced their calling time with increasing age (1 to 7 d). GC-MS analysis of hexane extracts of pheromone glands revealed that the titers of the five sex pheromone components were very low on the first night after eclosion, but increased sharply and peaked on the third night. Titers decreased greatly with increasing age (3 to 7 d), but from the onset of the scotophase to 4.5 h into scotophase, titers remained invariate. However, from 4.5 to 7 h, pheromone titers, especially that of the primary component Z7-12:Ac, increased sharply and peaked. After the peak, the titers decreased greatly at 8 h, and thereafter decreased gradually to the level before 4.5 h. The quantity of each component almost reached zero after lights-on.     

Diel periodicity and influence of age and mating on female sex pheromone titre in Estigmene acrea (Lep., Arctiidae)

Abstract:  Calling behaviour, diel periodicity, and effect of age and mating on female sex pheromone titre in Estigmene acrea (Drury) were studied under laboratory conditions. Forty-five per cent of females started calling during the first scotophase, but the highest number of calling females was observed during the second, third and fourth scotophases. Calling behaviour occurred from the third hour after dark until just before the end of the scotophase. However, females exhibited a bimodal pattern of calling with the first peak occurring between 4 and 6 h and a second peak at 10 h after the onset of scotophase. The mean onset of calling time differed significantly with age. Older females showed a tendency to call longer, but there was no significant difference. The amount of (Z,Z)-3,6-cis-9, 10-epoxyheneicosadiene in females was quantified from the first scotophase following emergence, until the fifth scotophase. Glands of 0-day-old females presented a higher content of pheromone compared with that found in glands of 1-, 2-, 3- and 4-day-old females. Pheromone titre was determined at 2-h intervals throughout the third scotophase and photophase. (Z,Z)-3,6-cis-9,10-epoxyheneicosadiene was found in the gland during the scotophase as well as the photophase. However, there was no consistent pattern of pheromone production throughout the scotophase or photophase. Mated females of E. acrea produced significantly less pheromone than virgin females.     

Sex pheromone titre in the glands of Spodoptera litura females: circadian rhythm and the effects of age and mating

The present study investigates the effects of age and mating status on the circadian variations of gland sex pheromone titre in female Spodoptera litura Fabricius. Similar to other nocturnal moths, S. litura females exhibit circadian variations of gland sex pheromone contents, with higher levels during scotophase and lower levels during photophase. The sex pheromone titre in the glands peaks during the first scotophase after eclosion and sharply declines afterwards. Higher pheromone contents during scotophase may facilitate female reproductive activities, and the negative relationship between pheromone titre and female calling is likely the result of pheromone release during female calling. Interestingly, the present study demonstrates that mated S. litura females have significantly higher sex pheromone titre in their pheromone glands (PGs) than virgin females. This finding contrasts with all previous studies of other insect species, in which mating generally reduces the sex pheromone titre in female PGs. In S. litura, mating and male accessory gland fluids can suppress female calling behaviours and re‐matings. These results suggest that the suppression of female calling behaviours by mating and male accessory gland fluids may significantly reduce the release of sex pheromones and thus result in higher sex pheromone titre in the PGs of mated females.     

Physiological Mechanisms of Pheromonostatic Responses: effects of Adrenergic Agonists and Antagonists on Moth (Helicoverpa armigera) Pheromone Biosynthesis

The adrenergic agonists octopamine, tyramine and clonidine inhibited the normal pheromonotropic action due to PBAN (pheromone biosynthesis activating neuropeptide) in incubations of intersegmental tissues that are situated between the 8th and 9th abdominal segments of the moth ovipositor tip. This inhibition was reversed in the presence of the adrenergic antagonists phentolamine, yohimbine and chlorpromazine. Incubations of 8th segments alone, which do not produce pheromone, resulted in elevated levels of intracellular cAMP in the presence of octopamine. The physiological significance of this phenomenon is unclear. However, clonidine (an alpha(2) selective agonist) did not duplicate octopamine stimulation of intracellular cAMP in 8th segment cultures. In intersegmental membrane cultures clonidine successfully duplicated the octopamine inhibition of both pheromone and intracellular cAMP production. The physiological significance of octopaminergic receptors mediating the inhibitory response of intersegments was investigated by experiments in vivo. When PBAN was injected into photophase females the normal pheromonotropic activity due to the injected PBAN dropped after 2h. In the presence of clonidine, normal peak stimulatory levels were never attained and a faster decline was observed. Clonidine also inhibited the pheromonotropic response of 24h-decapitated females to PBAN. Adrenergic antagonists successfully reversed the inhibitory effect of clonidine in decapitated females, but did not reverse the effect of clonidine in photophase females. In addition, when clonidine was injected into female moths during the scotophase normal peak pheromone titers were reduced although no effect on calling behavior was observed. Copyright 1997 Elsevier Science Ltd. All rights reserved     

Mechanisms of premating isolation between Helicoverpa armigera (Hübner) and Helicoverpa assulta (Guenée) (Lepidoptera: Noctuidae)

Helicoverpa armigera and Helicoverpa assulta are sympatric sibling species, and in the laboratory they can interbreed and produce viable offspring. To assess the contributions of temporal barriers and sexual barriers to premating isolation, we investigated both the temporal rhythms of calling behavior and pheromone titers of H. armigera and H. assulta females and the behavioral responses of males to conspecific and heterospecific calling females in a wind tunnel. Both H. armigera and H. assulta females called throughout the scotophase, and there was more calling during the second half of the scotophase than during the first half. Maximal pheromone titer and maximal calling activity in H. armigera synchronously occurred at the sixth hour into the scotophase, whereas, in H. assulta, the maximal pheromone titer occurred 2 h before the peak of calling. Pheromone blend ratios of the two species were opposite and, within each species, changes in the ratio within the scotophase and at different ages were relatively small. Males of both H. armigera and H. assulta responded strongly to their conspecific calling females in the wind tunnel and completed the whole courtship sequence. In contrast, they did not land and had no copulation attempts in response to heterospecific calling females. These results show that the two species do not have obvious temporal differences in calling behavior and pheromone production, and the specificity of sex pheromone blend emitted by females plays a key role in their premating isolation. In addition, we summarized the potential isolation mechanisms of H. armigera and H. assulta.     

Photoperiod cues and the modulatory action of octopamine and 5-hydroxytryptamine on locomotor and pheromone response in male gypsy moths,Lymantria dispar

Experiments were conducted to determine whether the biogenic amines octopamine (OA) and 5-hydroxytryptamine (5-HT) exert modulatory effects on pheromone responsiveness and random locomotor activity in male gypsy moths. When injected into males, OA significantly enhanced sensitivity to pheromone, while 5-HT enhanced general locomotor activity, results that were very similar to those previously shown for the cabbage looper. Maximal effect of the amines, however, was observed when injection occurred just prior to the onset of scotophase, rather than photophase, as we had originally hypothesized for this diurnally active insect. Male gypsy moths also displayed a prominent scotophase response, with sensitivity to pheromone greater in the scotphase compared with photophase, but with the level of random locomotor activity lower in scotophase than in photophase. The upwind flight behavior of males to a pheromone source in a wind tunnel, as well as the time spent at the source, were also significantly different in the two light regimes. Furthermore, when exposed to a 1 h scotophase (instead of the normal 8), or to continuous dark conditions, while males exhibited response to pheromone and locomotor activity during the same scotophase and photophase periods as observed in a 16:8 light : dark cycle, the levels of response, as well as qualitative aspects of the upwind flight behaviors in both periods were a function of the light intensity. Our combined results suggest that male gypsy moths display a bimodal rhythm of locomotor and pheromone response over the diel cycle, with light intensity and scotophase onset providing critical cues for the expression of behaviors, as well as the modulatory action of the amines. © 1992 Wiley-Liss, Inc.