Right-handed penises of the earwig Labidura riparia (Insecta, Dermaptera, Labiduridae): evolutionary relationships between structural and behavioral asymmetries

The number of penises vary in the insect suborder Forficulina (order Dermaptera; earwigs). Males of the families Diplatyidae, Pigidicranidae, Anisolabididae, Apachyidae, and Labiduridae have two penises (right and left), while those of the Spongipohridae, Chelisochidae, and Forficulidae have a single penis. The proposed phylogenetic relationships among these families suggest that the single‐penis families evolved from an ancestor possessing two penises. To date, examinations of double‐penis earwig species have found that only a single penis is used per single copulation. These diversities in structural and behavioral aspects of genitalia raises the following intriguing questions: How are the two penises used? Why did a penis degenerate in several earwig families, and which one was lost? To address these questions, structural and behavioral asymmetries were examined in detail for a representative species Labidura riparia (Labiduridae). Although there was no detectable morphological differentiation between the right and left penises, male L. riparia predominantly used the right one for insemination. This significant “right‐handedness” developed without any experience of mating and was also manifested in the resting postures of the two penises when not engaged in copulation. However, surgical ablation of the right penis did not influence the insemination capacity of males. In wild‐caught males, only about 10% were left‐handed; within this group, abnormalities were frequently observed in the right penis. These lines of evidence indicate that the left penis is merely a spare intromittent organ, which most L. riparia males are likely never to use. Additional observations of five species of single‐penis families revealed that the left penis degenerated in the common ancestor of this group. Considering the proposed sister relationship between the Labiduridae and the single‐penis families, it is possible that such behavioral asymmetries in penis' use, as observed in L. riparia, are parental to the evolutionary degeneration of the infrequently used left penis. J. Morphol., 2006. © 2006 Wiley‐Liss, Inc.     

Tissue clearing‐based method for unobstructed three‐dimensional imaging of mouse penis with subcellular resolution

Although mice are widely used to elucidate factors contributing to penile disorders and develop treatment options, quantification of tissue changes upon intervention is either limited to minuscule tissue volume (histology) or acquired with limited spatial resolution (MRI/CT). Thus, imaging method suitable for expeditious acquisition of the entire mouse penis with subcellular resolution is described that relies on both aqueous‐ (clear, unobstructed brain imaging cocktails and computational analysis) and solvent‐based (fluorescence‐preserving capability imaging of solvent‐cleared organs) tissue optical clearing (TOC). The combined TOC approach allows to image mouse penis innervation and vasculature with unprecedented detail and, for the first time, reveals the three‐dimensional structure of murine penis fibrocartilage.     

Resurrecting embryos of the tuatara,Sphenodon punctatus,to resolve vertebrate phallus evolution

The breadth of anatomical and functional diversity among amniote external genitalia has led to uncertainty about the evolutionary origins of the phallus. In several lineages, including the tuatara, Sphenodon punctatus, adults lack an intromittent phallus, raising the possibility that the amniote ancestor lacked external genitalia and reproduced using cloacal apposition. Accordingly, a phallus may have evolved multiple times in amniotes. However, similarities in development across amniote external genitalia suggest that the phallus may have a single evolutionary origin. To resolve the evolutionary history of amniote genitalia, we performed three-dimensional reconstruction of Victorian era tuatara embryos to look for embryological evidence of external genital initiation. Despite the absence of an intromittent phallus in adult tuataras, our observations show that tuatara embryos develop genital anlagen. This illustrates that there is a conserved developmental stage of external genital development among all amniotes and suggests a single evolutionary origin of amniote external genitalia.     

Comparative ultrastructure of the terminal portions of the male copulatory apparatus in Planorbidae (Gastropoda: Pulmonata)

Terminal portions of the male copulatory apparatus of Planorbis planorbis, Segmentina oelandica, and Anisus vortex were studied using whole-mount preparations, serial semi-thin sections, and transmission electron microscopy. In the latter species, stylet formation was investigated at several stages of postembryonic development. Organization of the penial distal portion in the species studied varies greatly. In P. planorbis, the distal end of the penis lacks developed papillae and is armed with a stylet built up of the covering epithelial cells of the penis proper. In A. vortex, the stylet is formed by the secretory activity of the middle cells of the distal portion of the penis. To the time of maturation, the cells encompassing the stylet are broken down exposing its solid chitinous structure and characteristic shape. In S. oelandica, the distal end of the penis bears the long probably flexible papilla with the characteristics of an internal ‘skeleton,’ organized as a line of connective tissue cells and a system of hydrocoelic cavities.     

Phylogenetic and taxonomic problems in freshwater Oligochaeta with special emphasis on chitinous structures in Tubificinae

Setae and chitinous penis sheaths are the main characters used to distinguish genera and species of the subfamily Tubificinae. Genital setae and penis sheaths are of functional importance to facilitate copulation. Similar structures in different genera may be homologous or products of parallel evolution (homoiologous). Form and dimensions of the penis sheaths of many tubificine species are very variable. Transspecific overlap of quantitative species characters can make the determination of specimens in the genus Limnodrilus difficult. The configuration of the distal ends of the penis sheaths is an important character to distinguish Limnodrilus species. The definition of the intraspecific variability of this morphological character is problematic.     

阴茎中脊囊肿的形态学观察

目的观察阴茎中脊囊肿临床病理及免疫组织化学特点，探讨其发生机理。方法3例阴茎中脊囊肿患切除标本进行常规HE染色及免疫组织化学染色。一抗分别为CK7，CK13，CK20，EMA，CEA，Vimentin，CgA，Syn)。结果所有3例患均以龟头形态异常就诊，囊肿均位于尿道外口旁。3例患囊肿被覆上皮均为尿道上皮及其化生上皮。免疫组织化学上皮细胞Keratin(CK7 ，CK13 ，CK20-)，EMA ，CEA ，Vimentin-，CgA-，Syn-。结论阴茎中脊囊肿可能起源于尿道外口旁外部导管来源的尿道上皮及其化生上皮，与胚生殖沟发育障碍有关。     

The role of female investment in a sexual arms race

Forced copulations are common among waterfowl species (Anatidae), a group with relatively large eggs and very precocial young. They can be viewed as the outcome of an overt sexual conflict, and an evolutionary arms race between the sexes. We examined the female choice hypothesis suggested, but not properly tested by Briskie and Montgomerie (1997) and Montgomerie and Briskie (2007), that penis size in male ducks might correlate with female investment in eggs, predicting that in species where females lay larger eggs, penis size might be larger, because females would be more reluctant to abandon their eggs if forced to copulate. A larger data set than in earlier studies enabled us to test that hypothesis in a comparative way. Our results compelled us to reject the female choice hypothesis since egg size is negatively correlated with penis length and the number of vaginal spirals, both being seen as adaptations to frequent forced copulations. The apparent trade-off between egg size and morphological defences (vaginal spirals) is strong particularly among monogamous species. Overall, we conclude that factors that set a lower limit for egg size constrain the morphological defences of females and the arms race between the sexes in waterfowl.     

Musculature of the penial complex: A new criterion in unravelling the phylogeny of Hygrophila (Gastropoda: Pulmonata)

The taxonomy of freshwater pulmonates (Hygrophila) has been in a fluid state warranting the search for new morphological criteria that may show congruence with molecular phylogenetic data. We examined the muscle arrangement in the penial complex (penis and penis sheath) of most major groups of freshwater pulmonates to explore to which extent the copulatory musculature can serve as a source of phylogenetic information for Hygrophila. The penises of Acroloxus lacustris (Acroloxidae), Radix auricularia (Lymnaeidae), and Physella acuta (Physidae) posses inner and outer layers of circular muscles and an intermediate layer of longitudinal muscles. The inner and outer muscle layers in the penis of Biomphalaria glabrata consist of circular muscles, but this species has two intermediate longitudinal layers separated by a lacunar space, which is crossed by radial and transverse fibers. The muscular wall of the penis of Planorbella duryi is composed of transverse and longitudinal fibers, with circular muscles as the outer layer. In Planorbidae, the penial musculature consists of inner and outer layers of longitudinal muscles and an intermediate layer of radial muscles. The penis sheath shows more variation in muscle patterns: its muscular wall has two layers in A. lacustris, P. acuta, and P. duryi, three layers in R. auricularia and Planorbinae and four layers in B. glabrata. To trace the evolution of the penial musculature, we mapped the muscle characters on a molecular phylogeny constructed from the concatenated 18S and mtCOI data set. The most convincing synapomorphies were found for Planorbinae (inner and outer penis layers of longitudinal muscles, three-layered wall of the penis sheath). A larger clade coinciding with Planorbidae is defined by the presence of radial muscles and two longitudinal layers in the penis. The comparative analysis of the penial musculature appears to be a promising tool in unraveling the phylogeny of Hygrophila.