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Although testosterone levels and muscle mass decline with age, many older men have serum testosterone level in the normal range, leading to speculation about whether older men are less sensitive to testosterone. We determined the responsiveness of androgen-dependent outcomes to graded testosterone doses in older men and compared it to that in young men. The participants in this randomized, double-blind trial were 60 ambulatory, healthy, older men, 60-75 yr of age, who had normal serum testosterone levels. Their responses to graded doses of testosterone were compared with previous data in 61 men, 19-35 yr old. The participants received a long-acting GnRH agonist to suppress endogenous testosterone production and 25, 50, 125, 300, or 600 mg testosterone enanthate weekly for 20 wk. Fat-free mass, fat mass, muscle strength, sexual function, mood, visuospatial cognition, hormone levels, and safety measures were evaluated before, during, and after treatment. Of 60 older men who were randomized, 52 completed the study. After adjusting for testosterone dose, changes in serum total testosterone (change, -6.8, -1.9, +16.1, +49.5, and +101.9 nmol/liter at 25, 50, 125, 300, and 600 mg/wk, respectively) and hemoglobin (change, -3.6, +9.9, +20.9, +12.6, and +29.4 g/liter at 25, 50, 125, 300, and 600 mg/wk, respectively) levels were dose-related in older men and significantly greater in older men than young men (each P

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The egg-laying apparatus consists of the uterus, the uterine muscles, the vulva, the vulval muscles, and a local neuropil formed by the egg-laying neurons (EggFIG 1). After fertilization, embryos pass from the spermatheca to the uterus, an epithelial egg chamber that links the two arms of the gonad. There, eggs develop to the approximately 30-cell stage (roughly 2.5 hr post-fertilization at 20C) before being expelled into the environment via the vulva, a passageway from the uterus to the ventral exterior. Under optimal conditions, an adult hermaphrodite will lay 4-10 eggs/hour. Egg-laying is facilitated by contraction of the sex muscles: the vulval muscles, which attach to the lips of the vulva, and the uterine muscles, which encircle the uterus. Muscle activity is regulated by motor neurons, in particular motor neurons VCn (VC1-6) and HSNL/R, which synapse onto each other and onto vulval muscle arms, forming a neuropil near the vulva. Tissues comprising the egg-laying apparatus arise from several different lineages (see ReproTABLE 1). As described below, the developing gonad and vulva act as organizing centers, recruiting cells from other regions to the midbody, coordinating cell patterning between different tissues, and directing axon guidance and synaptic patterning of the neurons.

Before the first fertilization event, the lumen of the uterus is narrow and blocked by a series of inwardly projecting fingers that extend from the uterine lumen wall (see EggFIG 5B and EggFIG 11B below). After passage of the first egg, the mature uterus retains a few inward septa that may derive from these earlier fingers. Both the developing and the mature uterine lumen have a continuous thickening or electron-dense layer (possibly a glycocalyx or surface coat; see EggFIG 11C). This is also apparent on the lumenal (apical) membrane, lining projecting fingers, and septa.

Vulval development spans roughly the same period as uterine development: L3 to late L4. For a summary figure of the stages of vulval development, please see EggFIG Sup1. Establishment of the vulva requires the local deformation of existing ventral structures such as the ventral nerve cord (VNC) (EggFIG 1) and ventral body wall muscles (EggFIG 3A), which are deflected laterally in this region to accommodate the vulva. Vulval development can be divided into two phases: (1) vulval cell patterning and generation (EggFIG 7 and EggFIG 8) and (2) vulval morphogenesis (EggFIG 9). The molecular and genetic mechanisms underlying these processes, particularly cell patterning, have been studied extensively and are described in the following reviews and papers and references therein (Greenwald, 1997; Kim, 1997; Eisenmann et al., 1998; Levitan and Greenwald, 1998; Hanna-Rose and Han, 2000; Shemer and Podbilewicz, 2003; Ceol and Horvitz, 2004; Sundaram, 2004).

As part of the process of joining vulval and uterine lumens, the AC creates a hole in the apex of the developing vulva (EggFIG 8). In L3, while Pnp cells are dividing, the ventral hypodermal basal lamina and gonadal basal lamina break down precisely at the site of contact with the AC. The basolateral portion of the AC crosses through this gap, attaches to, then inserts between the descendants of the primary-fated P6p lineage cells. This invasion is stimulated by a diffusible signal from the primary cells (Sherwood and Sternberg, 2003). Later, P6p terminal progeny fuse, forming a toroid (vulF) around the invading AC process. The AC is then removed by heterotypic fusion with the utse, leaving a channel in the apex of the vulva (Newman et al., 1996). When the utse membrane is ruptured by passage of the first egg, uterine and vulval lumens become continuous. During late L4, the vulval muscles attach to the vulval epithelial tube and to the body wall (see below). The tube then partially everts (turns inside out), generating the adult vulva in which the lumen is closed until vulval muscles contract (EggFIG 10) (Sulston and Horvitz, 1977; Sharma-Kishore et al., 1999).

The uterine (um1L/R, um2L/R) and vulval (vm1L/R, vm2L/R) muscles (EggFIG 1 and EggFIG 11A), collectively referred to as the sex muscles, are required for moving eggs through the uterus and vulva. Only 4 the 16 sex muscles receive direct inputs from the egg-laying neurons. The remaining sex muscles are electrically coupled, either directly or indirectly, to these innervated muscles (see EggFIG 13). This configuration may serve to coordinate uterine and vulval contraction. The sex muscles are classified as nonstriated muscles because they do not have the striated appearance (typified by body wall muscle) normally attributed to the presence of an ordered array of multiple sarcomeres (muscle contractile units; see Muscle System - Somatic Muscle). Vulval muscles have a single sarcomere that extends along the entire muscle length and attaches to a discrete zone in the body wall at one end and to the vulva at the other end (White, 1988). The uterine muscle myofilament network seems to be anchored to a thin basal lamina on the surface facing the uterus. In contrast to the vulval muscles, the attachment points are randomly arrayed and this distribution of dense bodies is similar to that seen in vertebrate smooth muscles (see Muscle System - Nonstriated). 041b061a72


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