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The testicle (from Latin testiculus, diminutive of testis, meaning "witness" of virility, plural testes) is the male gonad in animals. Like the ovaries to which they are homologous, testes are components of both the reproductive system and the endocrine system. The primary functions of the testes are to produce sperm (spermatogenesis) and to produce androgens, primarily testosterone. Both functions of the testicle are influenced by gonadotropic hormones produced by the anterior pituitary. Luteinizing hormone (LH) results in testosterone release. The presence of both testosterone and follicle-stimulating hormone (FSH) is needed to support spermatogenesis. It has also been shown in animal studies that if testes are exposed to either too high or too low levels of estrogens (such as estradiol; E2) spermatogenesis can be disrupted to such an extent that the animals become infertile. Anatomy and physiology External appearance Almost all healthy male vertebrates have two testes. In mammals, the testes are often contained within an extension of the abdomen called the scrotum. In mammals with external testes it is most common for one testicle to hang lower than the other. While the size of the testicle varies, it is estimated that 21.9% of men have their higher testicle being their left, while 27.3% of men have reported to have equally positioned testicles. This is due to differences in the vascular anatomical structure on the right and left sides. Internal structure Duct system Under a tough membranous shell, the tunica albuginea, the testis of amniotes and some teleost fish, contains very fine coiled tubes called seminiferous tubules. The tubules are lined with a layer of cells (germ cells) that from puberty into old age, develop into sperm cells (also known as spermatozoa or male gametes). The developing sperm travel through the seminiferous tubules to the rete testis located in the mediastinum testis, to the efferent ducts, and then to the epididymis where newly created sperm cells mature. The sperm move into the vas deferens, and are eventually expelled through the urethra and out of the urethral orifice through muscular contractions. Primary cell types Within the seminiferous tubules Here, germ cells develop into spermatogonia, spermatocytes, spermatids and spermatozoon through the process of spermatogenesis. The gametes contain DNA for fertilization of an ovum Sertoli cells - the true epithelium of the seminiferous epithelium, critical for the support of germ cell development into spermatozoa. Sertoli cells secrete inhibin. Peritubular myoid cells surround the seminiferous tubules. Between tubules (interstitial cells) Leydig cells - cells localized between seminiferous tubules that produce and secrete testosterone and other androgens important for sexual development and puberty, secondary sexual characteristics like facial hair, sexual behavior and libido, supporting spermatogenesis and erectile function. Testosterone also controls testicular volume. Also present are: Immature Leydig cells Interstitial macrophages and epithelial cells. Blood supply and lymphatic drainage Blood supply and lymphatic drainage of the testes and scrotum are distinct: The paired testicular arteries arise directly from the abdominal aorta and descend through the inguinal canal, while the scrotum and the rest of the external genitalia is supplied by the internal pudendal artery (itself a branch of the internal iliac artery). The testis has collateral blood supply from 1. the cremasteric artery (a branch of the inferior epigastric artery, which is a branch of the external iliac artery), and 2. the artery to the ductus deferens (a branch of the inferior vesical artery, which is a branch of the internal iliac artery). Therefore, if the testicular artery is ligated, e.g., during a Fowler-Stevens orchiopexy for a high undescended testis, the testis will usually survive on these other blood supplies. Lymphatic drainage of the testes follows the testicular arteries back to the paraaortic lymph nodes, while lymph from the scrotum drains to the inguinal lymph nodes. Layers Many anatomical features of the adult testis reflect its developmental origin in the abdomen. The layers of tissue enclosing each testicle are derived from the layers of the anterior abdominal wall. Notably, the cremasteric muscle arises from the internal oblique muscle. The blood–testis barrier Large molecules cannot pass from the blood into the lumen of a seminiferous tubule due to the presence of tight junctions between adjacent Sertoli cells. The spermatogonia are in the basal compartment (deep to the level of the tight junctions) and the more mature forms such as primary and secondary spermatocytes and spermatids are in the adluminal compartment. The function of the blood–testis barrier (red highlight in diagram above) may be to prevent an auto-immune reaction. Mature sperm (and their antigens) arise long after immune tolerance is established in infancy. Therefore, since sperm are antigenically different from self tissue, a male animal can react immunologically to his own sperm. In fact, he is capable of making antibodies against them. Injection of sperm antigens causes inflammation of the testis (auto-immune orchitis) and reduced fertility. Thus, the blood–testis barrier may reduce the likelihood that sperm proteins will induce an immune response, reducing fertility and so progeny. Temperature regulation The testes work best at temperatures slightly less than core body temperature. The spermatogenesis is less efficient at lower and higher temperatures. This is presumably why the testes are located outside the body. There are a number of mechanisms to maintain the testes at the optimum temperature. Cremasteric muscle The cremasteric muscle is part of the spermatic cord. When this muscle contracts, the cord is shortened and the testicle is moved closer up toward the body, which provides slightly more warmth to maintain optimal testicular temperature. When cooling is required, the cremasteric muscle relaxes and the testicle is lowered away from the warm body and is able to cool. It also occurs in response to stress (the testicles rise up toward the body in an effort to protect them in a fight). There are persistent reports that relaxation indicates approach of orgasm. There is a noticeable tendency to also retract during orgasm. The cremaster muscle can reflexively raise each testicle individually if properly triggered. This phenomenon is known as the cremasteric reflex. The testicles can also be lifted voluntarily using the pubococcygeus muscle, which partially activates related muscles.