长爪沙鼠阴茎头的组织学研究及五种鼠阴茎头的扫描电镜观察

本文对８例长爪沙鼠（Ｍｅｒｉｏｎｅｓｕｎｇｕｉｃｕｌａｔｕｓ）幼年、亚成年、成年和老年雄性个体的阴茎头组织进行连续切片。结果表明,阴茎头的外环层和阴茎骨近支由阴茎海绵体衍生而来;内环层、尿道小瓣、阴茎骨远支和侧支由尿道海绵体衍生而成。采用扫描电镜观察了长爪沙鼠、子午沙鼠（Ｍ．ｍｅｒｉｄｉａｎｕｓ）、大沙鼠（Ｒｈｏｍｂｏｍｙｓｏｐｉｍｕｓ）和甘肃绒鼠（Ｅｏｔｈｅｎｏｍｙｓｅｖａ）、黑腹绒鼠（Ｅ．ｍｅｌａｎｏｇａｓｔｅｒ）共５种１６例的阴茎头表皮棘,发现其形态、数量和分布有属、种间差异性和相对稳定性,沙鼠属（Ｍｅｒｉｏｎｅｓ）３种的表皮棘均呈牛角状的圆锥体,斜向排列成行;而绒鼠属（Ｅｏｔｈｅｎｏｍｙｓ）内甘肃绒鼠为球状表皮棘,黑腹绒鼠为牛角状,排列均不规则。因此,阴茎头表皮棘与阴茎骨、阴茎头软体结构均有分类学意义。     

鼩鼱科内6 个种的阴茎形态学

阴茎形态学特征是研究哺乳动物亲缘关系的有力工具,尤其是在近缘种之间。本试验选取鼩鼱科内的黑齿鼩鼱(Blarinella quadraticauda)、小纹背鼩鼱(Sorex bedfordiae)、大纹背鼩鼱(S. cylindricauda)、云南鼩鼱(S. excelsus)、四川短尾鼩(Anourosorex squamipes)和灰麝鼩(Crocidura attenuata)6 个种共31 号阴茎标本观察其阴茎形态学特征。研究结果表明:这6 个种的阴茎均缺乏阴茎骨;在6 个种的阴茎横切面上,尿道靠近阴茎腹侧面为阴茎海绵体包裹,没有单独的尿道海绵体存在,输精管由阴茎海绵体包裹,在阴茎末端形成两条输精管侧支;黑齿鼩鼱的阴茎具有独特的舌状结构,可作为鉴定黑齿鼩鼱的一个重要特征,鼩鼱属内的小纹背鼩鼱、大纹背鼩鼱和云南鼩鼱的阴茎头界限很难观察到,而黑齿鼩鼱、四川短尾鼩和灰麝鼩的阴茎头则具有明显的界限。     

Correlative 3D‐imaging of Pipistrellus penis micromorphology: Validating quantitative microCT images with undecalcified serial ground section histomorphology

Detailed knowledge of histomorphology is a prerequisite for the understanding of function, variation, and development. In bats, as in other mammals, penis and baculum morphology are important in species discrimination and phylogenetic studies. In this study, nondestructive 3D‐microtomographic (microCT, µCT) images of bacula and iodine‐stained penes of Pipistrellus pipistrellus were correlated with light microscopic images from undecalcified surface‐stained ground sections of three of these penes of P. pipistrellus (1 juvenile). The results were then compared with µCT‐images of bacula of P. pygmaeus, P. hanaki, and P. nathusii. The Y‐shaped baculum in all studied Pipistrellus species has a proximal base with two club‐shaped branches, a long slender shaft, and a forked distal tip. The branches contain a medullary cavity of variable size, which tapers into a central canal of variable length in the proximal baculum shaft. Both are surrounded by a lamellar and a woven bone layer and contain fatty marrow and blood vessels. The distal shaft consists of woven bone only, without a vascular canal. The proximal ends of the branches are connected with the tunica albuginea of the corpora cavernosa via entheses. In the penis shaft, the corpus spongiosum‐surrounded urethra lies in a ventral grove of the corpora cavernosa, and continues in the glans under the baculum. The glans penis predominantly comprises an enlarged corpus spongiosum, which surrounds urethra and baculum. In the 12 studied juvenile and subadult P. pipistrellus specimens the proximal branches of the baculum were shorter and without marrow cavity, while shaft and distal tip appeared already fully developed. The present combination with light microscopic images from one species enabled a more reliable interpretation of histomorphological structures in the µCT‐images from all four Pipistrellus species. J. Morphol. 276:695–706, 2015. © 2015 Wiley Periodicals, Inc.     

The penis of elephant shrews (Mammalia: Macroscelididae)

The penis of elephant shrews is very long, extending cranially to near the sternum where it emerges from a small caudally-pointing prepuce. The distal half of the penis lies free in a sheath and is termed the glans. In most species the corpus cavernosum is the main vascular body and extends to the tip of the penis while the corpus spongiosum around the urethra becomes much reduced distally, but in Rhynchocyon all vascular bodies are small and most of the penis consists of connective tissue. The distal tip of the penis is elaborated into various shapes, characteristic for each genus as currently defined. These penile features have more similarities with soricids and paenungulates than with archontans and lagomorphs but they are likely to be primitive characters and so should be interpreted cautiously.     

The functional morphology of penile erection: tissue designs for increasing and maintaining stiffness

Inflatable penises have evolved independently at least fourtimes in amniotes, specifically in mammals, turtles, squamates,and the archosaurs. Males in these lineages therefore sharethe functional problem of building a penis out of soft and flexibletissues that can increase its flexural stiffness and resistbending during copulation. Research on penile erectile tissuesin mammals and turtles shows that these two taxa have convergentlyevolved an axial orthogonal array of collagen fibers to reinforcethe penis during erection and copulation; in both lineages,the collagen fibers in the array are crimped and folded in theflaccid penis. Collagen fiber straightening during erectionincreases the stiffness of the tissue and allows changes inpenile radius that increase its second moment of area: bothof these changes increase the flexural stiffness of the penisas a whole. And once erect, axial orthogonal arrays have thehighest flexural stiffness of any fiber arrangement. The highdegree of anatomical convergence (to the level of microanatomicalfeatures) within mammals and turtles suggests that the stiffnessrequirements for copulation produce an extremely restrictiveselective regime in organisms that evolve inflatable penises.     

Axial orthogonal fiber reinforcement in the penis of the Nine-banded Armadillo (Dasypus novemcinctus)

Examination of histological sections from flaccid and artificially erected nine-banded armadillo (Dasypus novemcinctus) penises confirms that the mammalian corpus cavernosum is the first known biological hydrostat reinforced by collagen fibers arranged at 0° and 90° to its long axis. The morphology of this axial orthogonal fiber array affects the mechanical behavior of mammalian penises during erection and copulation. Specifically, the axial orthogonal array gives the erect penis a reproducible shape, maximum size and resistance to tensile, compressive, and bending forces. These features are more appropriate for the mechanical regime associated with copulation than those found in structures reinforced by crossed-helical fibers, although the axial orthogonal array also gives the corpus cavernosum a tendency to fail by kinking. Crimped collagen fibers in the flaccid array as well as three-dimensional folding of the wall in the flaccid corpus cavernosum allow the structure to expand during erection. J. Morphol. 233:249–255, 1997. © 1997 Wiley-Liss, Inc.     

The canine baculum: the structure and mechanical properties of an unusual bone

The baculum is an extraskeletal bone located in the penis of a few species in several orders of mammals such as carnivores, insectivores, rodents, bats and primates. This study aims to describe the structure, architecture and mechanical properties of the canine baculum. To this end canine bacula from castrated and uncastrated dogs were collected and examined by light microscopy, micro-computed tomography (microCT) scanning, histological staining, and mechanical testing. Their mineral density and mechanical properties were compared with those of a typical skeletal bone (the radius) in the same dog. Furthermore, a numerical model of a representative baculum was created and its mechanical performance analyzed using the finite element method, in order to try to elucidate its function. Examination of light microscopy images of transverse sections shows that the baculum consists of a typical sandwich structure, with two cortical plates separated, and joined, by loose cancellous bone. MicroCT scans reveal that the mineral density is lower in the baculum than in the radius, both in castrated as well as in uncastrated dogs, resulting in much lower stiffness. Castration was found to decrease the mineral density in both the baculum and the radius. The most likely function of the baculum of the dog is to stiffen the penis to assist intromission, and its much lower mineral density compared to that of the radius may be a mechanism designed to decrease the stiffness somewhat, and thus reduce the risk of fracture during copulation.     

Blood flow to the corpus cavernosum penis in the bull and goat buck during penile quiescence

Bulls and bucks were used to study the blood flow into and out of the corpus cavernosum penis (CCP) during the non-erect state of the penis. When contrast medium was injected into the dorsal artery of the penis in a bull cadaver and into surgically implanted catheters of anesthetized bucks, it flowed into the cavernous spaces of the CCP via the penetrating arteries. When contrast medium was injected directly into the CCP of the bull and buck there was no evidence of vascular exits along the body of the penis. The only venous outlets from the CCP were in the crus penis area. In the non-erect penis the vascular pressure within the CCP was 17.8 mm Hg higher in the bull and 8.3 mm Hg higher in the buck at the glans area than the crus area. During the non-erect state of the penis most of the blood flow to the CCP is from the dorsal artery of the penis via the penetrating arteries. The results showed that the pressure gradient provides the means for flow of blood from the glans area toward the base or crus penis area. This flow prevents platelet aggregation and clot formation and provides nutrition to local tissue during the long periods of non-erection.     

Baculum length and copulatory behaviour in carnivores and pinnipeds (Grand Order Ferae)

Relationships between baculum length, body weight and copulatory behaviour were examined in 66 species of carnivores and pinnipeds (Grand Order Ferae). Elongated bacula occur in most carnivore species of the families Ursidae, Canidae, Procyonidae and Mustelidae as well as in all pinnipeds studied. By contrast, members of the family Felidae have short bacula in relation to their body weights. Elongate bacula are found in carnivores and pinnipeds with a prolonged single intromission (PI) copulatory pattern. This finding agrees with results of a previous study of baculum length and PI copulatory patterns in primates. The enlarged baculum may serve to strengthen the penis and protect the urethra during prolonged intromissions. The distal pole of the baculum may also assist sperm transport since in some species it projects beyond the tip of the penis and probably contacts the female's os cervix during copulation. It is possible that stimulation of the female's genitalia by the baculum might also be important in mammals which are induced ovulators (e.g. Mustelidae). However, it is notable that elongation of the baculum has also occurred in some groups where females ovulate spontaneously (e.g. Canidae, Primates).     

Histomorphology of the glans penis in Vespertilionidae and Phyllostomidae species (Chiroptera,Mammalia)

The penises of bats are taxonomically distinctive in size and shape. In addition, they are variable in microscopic anatomy, indicating that histomorphological studies of copulatory organs of bats may help understanding their successful reproductive strategies. We studied adult males of 13 species of vespertilionid and phyllostomid bats. Both families exhibited the basic structure of the vascular penis of mammals: the hydrostatic elements of the corpora cavernosa and the corpus spongiosum surrounding the urethra, as well as accessory cavernous tissue. Variation in the position and amount of the tissues were observed in these families. Vespertilionid bats have a small glans penis with abundant accessory cavernous tissue on the prepuce and a highly variable baculum. The baculum varied in size and morphology, even among congeneric species, such as the three Lasiurus species and the two Myotis species. Phyllostomid species possess no bacula, but vascular structures are present to produce penile stiffening, particularly on the glans. Variation in the microscopic anatomy of the phyllostomid prepuce was observed, for example, Artibeus species had accessory cavernous tissue surrounded by a tunica albuginea, but Carollia perspicillata had two bundles of striated musculature and some adipose tissue; abundant pigments were present in the prepuce of most species.     

Penile paralysis and paraphimosis associated with reserpine administration in a stallion

Reserpine was administered to an 8-yr-old Thoroughbred stallion at a dosage of 5 mg subcutaneously (s.c.) every 2 wk for a 2-mo period to control unmanageable behavior. Reserpine produced a satisfactory calming effect that lasted for about 2 wk. After the last injection, the stallion developed penile paralysis and was unable to retract his penis, resulting in paraphimosis and attendant penile edema. The prolapsed penis was reduced and kept within the prepuce by placing a purse string suture in the preputial orifice. Phenylbutazone was given orally (1 gm) and the stallion was exercised for 20 to 30 min twice daily. This treatment was continued for 20 d with little improvement. The purse string retention suture cut through the skin 5 d after the stallion was discharged from the clinic. The penis was then supported against the abdominal wall and the stallion was exercised by hand for 30 min each day. The stallion was not used for breeding within 34 mo after the last injection of reserpine. A breeding soundness examination was performed approximately 3 yr after the initial injury. At this time the stallion's penis was noted to extend 5 to 8 cm from the prepuce when in a detumescent state. Although the stallion protruded his penis when exposed to a mare in estrus, a full rigid erection was never attained. Examination of the penis revealed partial engorgement of the corpus cavernosum penis and a 2.5-cm-wide dorsal semi-circumferential depression of the penile shaft approximately 10 to 12 cm proximal to the glans penis. The penile shaft and glans penis distal to this depression were cooler than the proximal portion of the penis. Semen collection was attempted, aided by manual insertion of the penis into the artificial vagina. When serving the artificial vagina, no "belling" of the glans penis was observed, although ejaculation occurred. Semen evaluation indicated normal spermatozoal motility and morphology parameters. The stallion was able to breed several mares with manual assistance to guide the penis into the vagina and one mare was diagnosed pregnant.     

Distribution of aromatase and sex steroid receptors in the baculum during the rat life cycle: effects of estrogen during the early development of the baculum

The baculum, also called os penis, plays an important role during copulation. However, the hormonal regulation of its development remains to be elucidated. To determine the direct involvement of sex steroids in the development of the baculum of rats, the distributions of androgen receptors (ARs), aromatase, and estrogen receptor alpha (ESR1) were observed immunohistochemically. On Postnatal Day 1, the rudiment of the baculum expressed ARs, aromatase, and ESR1. In the proximal segment of the baculum of neonatal rats, ARs were expressed in the parosteal layer but not in the periosteum or osteoblasts. Aromatase was expressed from the parosteal layer to the endosteum, particularly in the inner osteogenic layer. ESR1 was also abundantly expressed in almost all cells from the parosteal layer to the endosteum. ARs, aromatase, and ESR1 were all abundantly expressed during the neonatal period in the hyaline cartilage of the proximal segment and in fibrocartilage of the distal segment of the baculum. Expression in all the tissues was attenuated in an age-dependent manner and became quite weak at puberty. To determine the effect of estrogen on the growth of the baculum, the aromatase inhibitor 1,4,6-androstatrien-3,17-dione (ATD) was subcutaneously injected daily into pregnant rats from Days 19 to 23 of gestation and into pups on postnatal Days 1, 3, 5, 7, and 9. On Day 10, the length of the baculum in the ATD-treated rats was significantly shorter than that in the controls, although the body weight did not change. These findings suggest that not only androgen but also locally aromatized estrogen is involved in the early growth and development of the baculum.     

沟牙田鼠的形态特征及分类地位探讨

沟牙田鼠是分布于中国四川西部和甘肃南部的一种罕见小型啮齿动物,迄今标本收藏甚少,种、属地位存在异议,有人认为应并入田鼠属。我们根据四川九寨沟自然保护区采到的16号不同年龄、性别的沟牙田鼠,就其牙齿形态和成年雄性的阴茎结构与其它田鼠类作了比较,发现其成体和幼体在M³和M₃形态上有很大不同。其他重要区别包括：上门齿宽大;下门齿外露部分很短,下门齿总长仅及下颌长的77％左右,远比其他田鼠小;M齿环呈圆弧形或豆形亦很特殊;成年雄性的阴茎骨近支烧瓶状,远支基部膨大与田鼠类差别很大。结果表明：沟牙田鼠属是有别于田鼠属的有效属。     

Communication between the corpus cavernosum penis and the corpus spongiosum penis in a bull diagnosed by modified contrast cavernosography: A case report

Traditional methods of penile cavernosography failed to reveal a vascular defect or shunt from the corpus cavernosum penis of a 6-yr-old Brahman bull with a history of acquired failure of penile erection. When a tourniquet was applied to the penis and the contrast medium was introduced into the corpus cavernosum penis under pressure, however, a communication between the corpus cavernosum penis and corpus spongiosum penis was demonstrated. In a normal bull, cavernosography with the introduction of contrast medium under pressure produced a normal radiographic appearance. It is suggested that modification of accepted techniques of cavernosography to incorporate injection of contrast medium under pressure is useful in diagnosing this particular type of vascular abnormality, and may also aid in the detection of smaller vascular shunts from the corpus cavernosum penis to the superficial penile vasculature.     

The vasculature of the rat penis: a scanning electron microscopic and histologic study.

As a basis for understanding the mechanism of erection in an animal model frequently used in research in reproductive biology, the angioarchitecture of the penis of the rat has been described using scanning electron microscopy. Study of the penile vasculature of the rat indicates that the corpora cavernosa penis and the corpus spongiosum are independent erectile tissues, each with its own arterial and venous vessels. The large vascular spaces and abundant smooth muscle of the penile crura are compatible with its role in regulating blood flow to more distal penile tissues. Helicine arteries of the crura, but not the parent deep penile artery or arteries elsewhere, have muscular cushions in their walls. The venous drainage of the penile crura is via subtunical veins which are thought to be compressed during erection to elevate pressure within the penis. Large, paired cavernous veins drain the shaft of the penis. A unique method for inhibiting blood flow from the penis is indicated by the division of the cavernous veins into smaller channels prior to joining the subtunical venous plexus. Erectile tissue in the bifid origins of the corpus spongiosum has abundant cavernous muscle, while in the remainder of the corpus spongiosum little smooth muscle lines the cavernous spaces. The cavernous spaces on either side of the urethra coalesce to form vessels, each of which communicates with cavernous spaces in the glans. In addition, a bypass of the glans is effected by communication of these vessels directly with the deep dorsal vein. The apparent absence of muscular pads in vessels of the spongiosum, the relative paucity of cavernous smooth muscle, and the ample venous drainage provided by the deep dorsal vein may account for the lack of a venous occlusive mechanism similar to that of the corpora cavernosa penis.     

Neuropeptide Y in the human male genital tract

Neuropeptide Y (NPY) was found in high concentrations in the male genital tract. NPY levels were highest in the seminal vesicles, prostate, corpus cavernosum and vas deferens, where large numbers of immunoreactive nerve fibres were detected. Considerable quantities were also found in the epididymis and spongiosum. Lower concentrations were found in the glans penis, testis and foreskin. The presence of a large number of nerves containing NPY suggest that this active neuropeptide may play a role in control of genital function.     

Immunohistochemical analysis of smooth muscle cells and volumetric density of the elastic system fibers of wild boar (Sus scrofa) penis

The purpose of the present study was to verify the smooth muscle cell distribution and elastic system fibers volumetric density (Vv) in the corpus spongiosum and corpus cavernosum of the wild boar penis. Adult wild boars (n=13) were used. The penile mid shaft fragments were fixed with 4% phosphate buffered formalin solution and/or Bouin's liquid during 24-48 h, and processed using standard histological techniques. The sections were stained with Weigert's Resorcin-Fucsin with previous oxidation. The elastic system fibers Vv was determined in 25 random fields of each fragment using M42 test system. For immunohistochemical analysis, monoclonal anti-alpha actin smooth muscle was used. The histochemical methods detected elastic system fibers in both corpus spongiosum and corpus cavernosum of all animals. The elastic fibers Vv average was 36.6%+/-0.9 for corpus spongiosum and 11.7%+/-0.5 for corpus cavernosum. Through immunocitochemistry, a small quantity of smooth muscle cells was visualized in intimate relationship with blood vessels wall. The great amount of elastic fibers and the smooth muscle cell distribution beneath the endothelium suggest that these fibers may have an important role in penile erection process in the penis of wild boars.     

Modifications of Erectile Tissue Components in the Penis during the Fetal Period

Background

The penile erectile tissue has a complex microscopic anatomy with important functions in the mechanism of penile erection. The knowledge of such structures is necessary for understanding the normal physiology of the adult penis. Therefore, it is important to know the changes of these penile structures during fetal development. This study aims to analyze the development of the main components of the erectile tissue, such as collagen, smooth muscle fibers and elastic system fibers, in human fetuses.

Methodology/Principal Findings

We studied the penises of 56 human fetuses aged 13 to 36 weeks post-conception (WPC). We used histochemical and immunohistochemical staining, as well as morphometric techniques to analyze the collagen, smooth muscle fibers and elastic system fibers in the corpus cavernosum and in the corpus spongiosum. These elements were identified and quantified as percentage by using the Image J software (NIH, Bethesda, USA). From 13 to 36 WPC, in the corpus cavernosum, the amount of collagen, smooth muscle fibers and elastic system fibers varied from 19.88% to 36.60%, from 4.39% to 29.76% and from 1.91% to 8.92%, respectively. In the corpus spongiosum, the amount of collagen, smooth muscle fibers and elastic system fibers varied from 34.65% to 45.89%, from 0.60% to 11.90% and from 3.22% to 11.93%, respectively.

Conclusions

We found strong correlation between the elements analyzed with fetal age, both in corpus cavernosum and corpus spongiosum. The growth rate of these elements was more intense during the second trimester (13 to 24 WPC) of gestation, both in corpus cavernosum and in corpus spongiosum. There is greater proportional amount of collagen in the corpus spongiosum than in corpus cavernosum during all fetal period. In the corpus spongiosum, there is about four times more collagen than smooth muscle fibers and elastic system fibers, during all fetal period studied.     

中国13种鼠科啮齿动物的阴茎形态学及其分类学意义的探讨

本文对我国所产13种鼠科啮齿类进行了阴茎形态学的比较研究(包括阴茎骨和软体结构),探讨了在鼠科的家鼠属(Rattus)、姬鼠属(Apodemus)、小家鼠属(Mus)和巢鼠属(Micromys)4个属间以及同属不同种之间的差异。结果表明,阴茎的形态结构具有属和种的特异性和稳定性,可以为分类学的研究提供重要依据。     

Fellatio by Fruit Bats Prolongs Copulation Time

Oral sex is widely used in human foreplay, but rarely documented in other animals. Fellatio has been recorded in bonobos Pan paniscus, but even then functions largely as play behaviour among juvenile males. The short-nosed fruit bat Cynopterus sphinx exhibits resource defence polygyny and one sexually active male often roosts with groups of females in tents made from leaves. Female bats often lick their mate''s penis during dorsoventral copulation. The female lowers her head to lick the shaft or the base of the male''s penis but does not lick the glans penis which has already penetrated the vagina. Males never withdrew their penis when it was licked by the mating partner. A positive relationship exists between the length of time that the female licked the male''s penis during copulation and the duration of copulation. Furthermore, mating pairs spent significantly more time in copulation if the female licked her mate''s penis than if fellatio was absent. Males also show postcopulatory genital grooming after intromission. At present, we do not know why genital licking occurs, and we present four non-mutually exclusive hypotheses that may explain the function of fellatio in C. sphinx.