Copulation in spiders: experimental evidence for fatigue effects and bilateral control of palpal insertions.

Male lycosid spiders were prevented from using one palp during the first third of copulation by a sling that we then cut to free the palp. Initial insertion durations of the treated palp were significantly shorter than those shown concurrently by the much-used normal palp; they showed temporal patterning unaffected by use of the normal palp. Data indicate that there are relatively independent control mechanisms for each palp and support the hypothesis that fatigue causes the increased duration of successive palpal insertions. Our data also suggest that normal matings terminate due to maximal use of either palp by the male.     

Copulation in the lycosid spider (Lycosa rabida walckenaer): A quantitative study

An event recorder was used to obtain information on the temporal patterning of palpal insertions and palpal moistening in wolf spider matings. The series of insertions of each palp was characterized by a linear increase in insertion duration. The rate of change of insertion duration was fairly constant for the species. Asymmetry in the mechanism underlying palpal insertion was indicated by differences between the palps of some males in mean insertion duration and repetition rate. Termination of mating was initiated by the male's decreasing activity. Defensive behaviours described for the first time in spiders included spine erection in both sexes during mounting and ‘cheliceral pinching’ by the male when dismounting.     

星豹蛛求偶和交配行为

以星豹蛛(Pardosa astrigera)为研究对象,在室内对其求偶和交配行为进行了描述。雄蛛"俯卧撑"式动作(push-up)在求偶中具重要作用。交配初期,两侧触肢交替插入;随交配进行,单侧触肢连续插入3~5次后才换另一侧触肢插入,触肢每插入一次,基血囊膨大多次。完整交配一次雄蛛触肢器平均插入次数为29·625。交配前求偶时间、交配持续时间和有效交配时间分别平均为6min、32min25s和11min11s。星豹蛛雄蛛可进行多次交配,而雌蛛一般为单次交配。雌蛛交配状态(是否已经交配)影响其同类相食行为,已交配雌蛛对雄蛛同类相食率显著高于未交配雌蛛对雄蛛同类相食率。     

桑天牛头部附器感器的扫描电镜观察

利用扫描电镜观测桑天牛Apriona germari(Hope)成虫的触角、下颚须和下唇须上感器的分布及超微结构。结果表明,雌、雄桑天牛触角上共存在6种感器,即毛形感器、锥形感器、刺形感器、芽形感器、鳞形感器和棒形感器,其中锥形感器分5种亚型,刺形感器分2种亚型,且部分感器在雌雄成虫触角上的分布模式及数量存在差异,如棒形感器及角锥形感器丛模式仅在雄虫触角上发现,而雌虫触角上的细锥形感器和耳锥形感器多于雄虫,芽形感器少于雄虫;下颚须和下唇须存在5种感器,即毛形感器、刺形感器、末梢锥形感器、钟形感器和隙缝感器,各种感器在雌、雄天牛下颚须和下唇须上的分布和数量无明显的区别。     

浅斑近微蛛体形大小对交配行为的影响

理论和实验表明体形大小是许多动物择偶的重要信号。体形大小与产卵大小、最终产卵数量、后代适合度等都有着密切的关系,因此动物利用体形大小为信号来择偶可以获得高的生殖利益。目前研究交配行为和体形大小的关系的文献较少,特别在微蛛亚科。本文以浅斑近微蛛为实验材料,在室内研究了体形大小对交配行为的影响。测量了体重,体长,头胸甲与腹部的长和宽,以头胸甲的面积(头胸甲的长与宽的乘积)为变量来表示体形大小。以雄蛛的行为为依据,将其交配行为划分为4个时期:潜伏期、求偶期、插入期和纳精期,并记录了各个时期的持续时间。通过分析,发现了一种微蛛新的交配模式,即可将浅斑近微蛛的插入期分为三个阶段:首先是大约25min长插入期,接着是一个大约15min的中度插入期,最后是一个大约5min的短插入期。结果表明,在本试验条件下,浅斑近微蛛的体形大小对其交配行为的潜伏期、求偶期、插入期和纳精期的持续时间都没有影响,对插入期的3个阶段的长短也没有影响。     

Female sexual receptivity after partial copulations in the wolf spider (Schizocosa malitiosa)

Mated females of the wolf spider Schizocosa malitiosa are frequently refractory to further copulations. Copulation consists of two successive behavioural patterns: pattern I (PI) and pattern II (PII). During PI males make multiple consecutive insertions with each palp, and in PII males alternate the use of palps after each insertion until dismounting. As both patterns are inseminatory, another function – such as generating female reluctance – is suggested for this complex behaviour. Here we test experimentally whether female spiders mated only with PI, or only with PII, are reluctant to re-mate. Each copulating male was interrupted immediately after the end of PI (phase A), and the same male was instantly exposed to a second virgin female. After re-mounting, the male initially performed a brief recapitulation of PI followed by PII (both together considered as phase B). For testing female sexual reluctance, females were exposed to a second male 3 days after the partial mating, but whenever the mounting occurred it was interrupted. Females were raised and progeny was counted. Phase-A-mated females were more reluctant to re-mating than phase-B-mated females, but their respective progeny was similar. We suggest that female reluctance is caused by sperm-associated substances transferred during PI.     

Carbon isotope signatures reveal that diet is related to the relative sizes of the gills and palps in bivalves

In marine bivalves, the relative sizes of the gills and palps appear to be a useful functional trait that reflect feeding mode, i.e. suspension feeders have relatively larger gills than palps for pumping, whereas deposit feeders have relatively larger palps than gills for sorting. Also, within a species, the relative sizes of the gills and palps are related to changes in local food conditions. However, there is still no firm evidence showing that differences in the relative gill and palp sizes between species are related to diet selection. Based on the knowledge that carbon and nitrogen isotope signatures of an animals tissues reflect past diet, we compared the relative gill and palp sizes of bivalves from Roebuck Bay, northwestern Australia with their carbon and nitrogen isotope signatures. The carbon isotope signatures distinguished clear differences in diet between bivalves along a gradient from suspension to deposit feeding, and strikingly this pattern was closely followed by the relative sizes of the gills and palps of the bivalves. This study confirms that relative gill and palp sizes in bivalves are a functional trait that can be used to compare resource use between species. Furthermore, these data may suggest that morphospace occupation, as determined by relative gill and palp sizes of bivalves, could reflect a gradient of resource use between species.     

Chemo-discriminatory neurones in the sub-oesophageal ganglion of Locusta migratoria

The sub-oesophageal ganglion of Locusta migratoria was searched for neurones responsive to stimulation of the maxillary palps using intracellular recording techniques. Two plant stimuli were used: wheat, a host plant and cabbage, an unacceptable non-host plant. The stimuli were presented to the palp as both intact leaf tissue and as droplets of aqueous solutions of plant extracts. All the sampled neurones that responded to stimulation of the palp also responded to simple mechanical stimulation. However, 25% of the neurones exhibited consistent differences in response to the two plants when presented as both leaf tissue and droplets, strongly suggesting that they also received a chemosensory input. These differential responses most commonly took the form of differences in the duration of cell activity and/or variation in the latency of the onset of response. The receptive fields of differentially responding neurones were confined to the maxillary palp, or at most to the maxillary and labial palps.     

Multimodal chemosensory integration through the maxillary palp in Drosophila

Drosophila melanogaster has an olfactory organ called the maxillary palp. It is smaller and numerically simpler than the antenna, and its specific role in behavior has long been unclear. Because of its proximity to the mouthparts, I explored the possibility of a role in taste behavior. Maxillary palp was tuned to mediate odor-induced taste enhancement: a sucrose solution was more appealing when simultaneously presented with the odorant 4-methylphenol. The same result was observed with other odors that stimulate other types of olfactory receptor neuron in the maxillary palp. When an antennal olfactory receptor was genetically introduced in the maxillary palp, the fly interpreted a new odor as a sweet-enhancing smell. These results all point to taste enhancement as a function of the maxillary palp. It also opens the door for studying integration of multiple senses in a model organism.     

Regulation of Drosophila neural development by a putative secreted protein

The Drosophila strawberry (sty) locus was isolated by P-element insertion mutagenesis in a screen for mutations affecting eye development. Analysis of the mutant phenotype and the putative expression pattern of the sty gene suggested that it has multiple functions. Mutations in the sty gene lead to irregular spacing of ommatidia, an increase in the number of photoreceptor cells, as well as abnormal axonal projections to the lamina and disrupted structure of the optic lobes in the adult fly. The sty mutation also causes abnormal head involution, a change in a number of sensilla in the antennomaxillary complex in the embryonic stage and abnormal morphogenesis of the maxillary palp and wings in later stages. We examined the presumptive expression of the sty gene during development by histochemical staining for lacZ expression from enhancer trap elements inserted within the sty gene. During embryogenesis, expression of lacZ showed a segmental pattern in the ectoderm and in the nervous system. In the eye imaginal discs, lacZ was expressed in photoreceptor cells beginning a few rows posterior to the morphogenetic furrow. The lacZ was also expressed in the wing disc. In the adult, lacZ was expressed in the retina and lamina. We cloned the sty gene by P-element tagging and found that it encodes a putative secreted protein containing a cysteine-rich region similar to the epidermal growth factor (EGF) repeat. On the basis of the loss of functional phenotype, the expression pattern and the predicted structure of its product, we propose that sty encodes a diffusible protein acting as a signal involved in lateral inhibition within the developing nervous system and also as a factor involved either directly or indirectly in axonal guidance and optic lobe development.     

Sexual dimorphism of adult labial palps of the peach fruit moth Carposina sasakii Matsumura (Lepidoptera: Carposinidae) with notes on their sensilla

The labial palps and their sensilla of the peach fruit moth Carposina sasakii Matsumura, a serious pest of pome fruits in eastern Asia, were investigated using light microscopy and scanning and transmission electron microscopy. The labial palps are three‐segmented and exhibit distinct sexual dimorphism in length: much longer in the female than in the male. Four types of sensilla were found on the labial palps: campaniform, squamiform, chaetic and flattened sensilla. The campaniform sensilla are present on the first segment alone. The squamiform sensilla are located on all the three segments. The chaetic sensilla are mainly present on the third segment. The flattened sensilla are grooved with wall pores and situated in a labial palp‐pit organ, which is located at the apex of the distal segment of the labial palp. The sexual dimorphism of the labial palp and the labial palp‐pit organ was briefly discussed.     

Morphology of anterior regeneration in two spionid polychaete species: implications for feeding efficiency

Abstract. In marine soft‐sediment habitats, tissue loss by infuanal invertebrates can provide significant energetic input to higher trophic levels, have substantial impacts on individual behavior, growth, and fecundity, and resulting changes in bioturbation rates can secondarily affect community dynamics. The degree to which a community is affected by such sublethal predation depends in part on whether injured individuals can regenerate and on the speed at which they do so. Previously, we demonstrated differences in the rate of anterior segment and palp regeneration by the spionid polychaetes Pygospio elegans and Dipolydora quadrilobata. The current study examined the morphology of the anterior segment and palp regeneration in these species using scanning electron microscopy (SEM) and indirect immunohistochemistry with confocal laser scanning microscopy at 3, 6, 9, and 12 d post‐ablation. Antibodies for acetylated α‐tubulin and the neural tetrapeptide FMRFamide were used to label the regenerating nervous system. SEM revealed that the morphology of anterior tissue regeneration was similar for both species, but the ciliated food groove tended to form sooner on palps in P. elegans than on those of D. quadrilobata. In both species, palp regeneration and ciliated food groove formation were faster when only palps were removed. A shortened ciliated food groove is likely to reduce particle contact and transport efficiency in suspension and deposit feeding. Regenerating palp nerves were initially visible at 3 d following ablation of palps only, but at 6 d following ablation of five anterior segments. Following ablation of anterior segments, the regenerating nervous system was largely complete by 9 d, nuchal organs were innervated by 6 d, and processes of palp sensory cells were visible at 12 d. Contact chemoreception by sensory cells on the palps may be diminished during the early stages of regeneration, but chemoreception of waterborne cues via the nuchal organ should not.     

The maxillary palp of Drosophila : ultrastructure and physiology depends on the lozenge gene

The ultrastructure and physiology of the maxillary palp of Drosophila melanogaster have been studied in wild-type and lozenge mutants. Olfactory physiology in the maxillary palp is shown to depend upon the lozenge(lz) gene. Reduced response amplitudes were recorded for all odorants tested, and the physiological defect was shown to map to the lz locus. The structure of the maxillary palp sensilla is described by scanning electron microscopy (SEM) at high magnification, initially in the wild-type. A linear arrangement of pores, connected by furrows, was found in one class of sensilla, the basiconic sensilla. In the lz ³ mutant, morphological alterations in the basiconic sensilla and duplications of sensilla are documented by SEM. The correlation of structural abnormalities in the lz sensilla and physiological abnormalities in odorant response are consistent with an olfactory role for the basiconic sensilla of the maxillary palp. Accepted: 10 September 1996     

Retinoic acid differently affects the formation of palps and surrounding neurons in the ascidian tadpole

The anterior-most surface of the ascidian tadpole larvae is composed of specialized complex structures, including adhesive organs (palps) and the surrounding sensory neurons (RTENs) connected to neurons inside the palps. These are derived from a-line blastomeres by inductive effects from A-line blastomeres. The induction is reported to coordinate the expression of homeobox genes in the anterior epidermis, which can be affected by all- trans retinoic acid (RA). RA treatment also results in failure of the morphological formation of palps. Here we first isolated a gene intensely expressed in the cells of the anterior structure from the time of their lineage restriction, and then found that the RA treatment did not affect the specific gene expression in the presumptive palp cells but did affect that in the RTENs. These results suggest that the palp formation involves at least two different processes, a RA-insensitive cell-type specification process and a RA-sensitive morphogenetic process. RA treatment also affects the morphogenetic process of the palp formation and also disturbs the precise patterning of the surrounding epidermis, which may contribute to the regulation of RTEN development.     

Analysis of Drosophila proboscipedia mutant alleles.

Proboscipedia (PB) is a HOX protein required for adult maxillary palp and proboscis formation. To identify domains of PB important for function, 21 pb point mutant alleles were sequenced. Twelve pb alleles had DNA sequence changes that encode an altered PB protein product. The DNA sequence changes of these 12 alleles fell into 2 categories: missense alleles that effect the PB homeodomain (HD), and nonsense or frameshift alleles that result in C-terminal truncations of the PB protein. The phenotypic analysis of the pb homeobox missense alleles suggests that the PB HD is required for maxillary palp and proboscis development and pb - Sex combs reduced (Scr) genetic interaction. The phenotypic analysis of the pb nonsense or frameshift alleles suggests that the C-terminus is an important region required for maxillary palp and proboscis development and pb-Scr genetic interaction. PB and SCR do not interact directly with one another in a co-immunoprecipitation assay and in a yeast two-hybrid analysis, which suggests the pb-Scr genetic interaction is not mediated by a direct interaction between PB and SCR.     

Spontaneous male death during copulation in an orb-weaving spider

Males of some cannibalistic species of spiders and insects appear to sacrifice themselves by allowing the female to eat them, and the adaptive significance of such drastic terminal reproductive investment has recently been demonstrated for a spider. Typically, the female has to kill the male, but it has been suggested that males of some species in the cannibalistic orb-weaving spider genus Argiope may die in copula without female 'collaboration'. Here, we provide the first experimental evidence to our knowledge of programmed sudden death after onset of copulation in males of the spider Argiope aurantia. Our observations reveal that males invariably die during the insertion of their second pedipalp, regardless of whether they mate with newly moulted, defenceless females or with older mature females that often attack them. We determined experimentally that the death of males is triggered immediately upon insertion of the second palp, when males become unresponsive, and heartbeat ceases within minutes of insertion. We discuss the possible adaptive significance of programmed death during copulation, and argue that male death has evolved in a context other than sexual cannibalism.     

Maxillary palp glomeruli and ipsilateral projections in the antennal lobe of<Emphasis Type="Italic">Drosophila melanogaster</Emphasis>

The antennal lobe was examined by Golgi-silver impregnation to differentiate the glomeruli depending on the source and types of inputs. Thirty-five of the 43 ‘identified’ olfactory glomeruli were Golgi-silver impregnated in the present study. Seven glomeruli compared to three, reported previously, were found to be targets of maxillary palp chemosensory neurons. These include glomeruli VA3, VC2, VM5, VA7m/VA7l of the ventral antennal lobe and DC2, DC3, DM5 of the dorsal antennal lobe. The number of glomeruli receiving the maxillary palp sensory projections tallies with the number ofDrosophila olfactory receptors (seven) reported to be expressed exclusively in the maxillary palp. Twenty-eight Golgi-impregnated glomeruli were found to receive input from the antennal nerve. The ratio of glomeruli serving the maxillary palp to those serving the antenna (∼1:5) matches with the ratio ofDrosophila olfactory receptors expressed in these two olfactory organs respectively. In addition to glomerulus V, glomeruli VP1-3, VL1, VL2a/2p and VC3m/3l were found to receive ipsilateral projections. Thus, additional ipsilateral glomeruli have been identified.     

Olfactory physiology in the Drosophila antenna and maxillary palp: acj6 distinguishes two classes of odorant pathways.

This article provides characterization of the electrical response to odorants in the Drosophila antenna and provides physiological evidence that a second organ, the maxillary palp, also has olfactory function in Drosophila. The acj6 mutation, previously isolated by virtue of defective olfactory behavior, affects olfactory physiology in the maxillary palp as well as in the antenna. Interestingly, abnormal chemosensory jump 6 (acj6) reduces response in the maxillary palp to all odorants tested except benzaldehyde (odor of almond), as if response to benzaldehyde is mediated through a different type of odorant pathway from the other odorants. In other experiments, different parts of the antenna are shown to differ with respect to odorant sensitivity. Evidence is also provided that antennal response to odorants varies with age, and that odorants differ in their age dependence.     

Host-handling behavior of the parasitoid,Diglyphus minoeus [Hym.: Eulophidae], parasitizing the honeysuckle leaf-miner,Phytomyza lonicerae [Dipt.: Agromyzidae]

The host-handling behavior of individual female waspsDiglyphys minoeus which attack the larvae of the honeysuckle leaf-miner,Phytomyza lonicerae, include 4 types of behavior: probing, ovipositor insertion, host-feeding, and resting. Ovipositor insertion may either be injection of venom, probing of the host by the sensilla on the ovipositor's tip or egg laying. Three types of attacks were distinguished: oviposition attack for unparasitized larvae, host-feeding attack for unparasitized host larvae and host rejection for previously parasitized host larvae. Oviposition attack was characterized by frequent alternation between probing and ovipositor insertion, long duration of ovipositor insertion and resting, and the long duration of host-handling. Resting behavior is thought to protect the progeny against superparasitism, host-feeding by other wasps or hosts' recovery from paralysis. Host-feeding attack was characterized by frequent alternation between host-feeding and ovipositor insertion and long duration of host-feeding. Host rejection was composed mainly of probing and ovipositor insertion and short handling time.      

Acoustic interneuron responses compared in certain hawk moths.

A comparison is made of the interneuron responses to acoustic stimuli delivered under comparable conditions in hawk moths of six choerocampine species. There are marked interspecific differences in the duration of the afterdischarge and also in the pattern of the spike response train. The possible adaptive significance and the neurophysiological basis for these differences are discussed. Two non-choerocampine but mutually related hawk moth species show simpler patterns of interneuron response to acoustic stimulation. One of these moths appears to be able to detect the cries of bats flying nearby. However, the reception of sound in these two species does not involve the palp and pilifer, as it does in the choerocampines. Their auditory mechanism lies in the cranial-cervical region but it has not been identified. These analogous means of detecting ultrasonic pulses are discussed in relation to the sensitivity of comparable interneurons to vibrational (not airborne) sensitivity in other non-choerocampine hawk moths and in insects in general.