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Are consumers at risk when eating meat from animals that have been treated with growth-promoting hormones? Science, Technology and Medicine open access publisher.Publish, read and share novel research. E Rajakoski, The ovarian follicular system in sexually mature heifers with special reference to seasonal, cyclical, end left-right variations.Acta Endocrinol Suppl (Copenh). P Lamothe, D Bousquet, P Guay, Cyclic variation of F prostaglandins in the uterine fluids of the cow.J Reprod Fertil. Prunus cerasoides also called the Wild Himalayan Cherry is a deciduous tree found in East Asia.
Cultivation and propagationThe tree thrives in well-drained and moisture-retentive loamy soil. Basic vehicle dynamics and integration with the driver have also been examined, resulting in new componentry and creating a Human-Machine Interface (HMI) never seen before in the marketplace.
Taking these factors into account, Nissan created the all-new Nissan GT-R, which offers advanced high performance for secure and enjoyable driving by minimizing the effects of climatic, road condition or driving technique limitations.
6-speed Dual Clutch Transmission with three driver-selectable modes: Normal (for maximum smoothness and efficiency), Snow (for gentler starting and shifting on slippery surfaces), and R mode (for maximum performance with fastest shifts).
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Two-piece floating-rotor 15-inch front and rear discs with diamond-pattern internal ventilation. Driver-configurable performance monitor, developed with Sony  Polyphony, with graphical readouts of vehicle data and driving data displayed on a total of 11 screens.
7-inch WVGA high-resolution color-LCD display for audio, navigation and performance monitor. Growth hormone (GH) is a peptide hormone that stimulates growth and cell reproduction in humans and other animals. This hormone is used clinically to treat children's growth disorders and adult growth hormone deficiency. Peptides released by neurosecretory nuclei of the hypothalamus (Growth hormone releasing hormone and somatostatin) into the portal venous blood surrounding the pituitary are the major controllers of GH secretion by the somatotropes. In addition to control by endogenous and stimulous processes, a number of foreign compounds (xenobiotics) are known to influence GH secretion and function. GH can be used to treat conditions which produce short stature but are not related to deficiencies in GH, though results are not as dramatic when compared to short stature solely due to deficiency of GH. GH treatment improves muscle strength and slightly reduces body fat in Prader-Willi syndrome, which are significant concerns beyond the need to increase height.
Claims for GH as an anti-aging treatment date back to 1990 when the New England Journal of Medicine published a study where GH was used to treat 12 men over 60. A Stanford University School of Medicine survey of clinical studies on the subject published in early 2007 showed that the application of GH on healthy elderly patients increased muscle by about 2 kg and decreased body fat by the same amount  However, these were the only positive effects from taking GH. F Roche, Control and regulation of folliculogenesis--a symposium in perspective.Rev Reprod. Pattern of Plasma Luteinizing Hormone in the Cyclic Cow: Dependence upon the Period of the Cycle. Current concepts of the roles of follicle stimulating hormone and luteinizing hormone in folliculogenesis.
Evolution of the diameters of the largest healthy and atretic follicles during the human menstrual cycle. Selection of the Dominant Follicle in Cattle: Role of Two-Way Functional Coupling Between Follicle-Stimulating Hormone and the Follicles. Expression of follicle-stimulating hormone and luteinizing hormone receptor messenger ribonucleic acids in bovine follicles during the first follicular wave. Changes in messenger ribonucleic acid encoding luteinizing hormone receptor, cytochrome P450-side chain cleavage, and aromatase are associated with recruitment and selection of bovine ovarian follicles.
Gene expression for LH receptor, 17?-hydroxylase and StAR in the theca interna of preantral and early antral follicles in the bovine ovary. Development of Nonovulatory Antral Follicles in Heifers: Changes in Steroids in Follicular Fluid and Receptors for Gonadotr opins.
Development of Antral Follicles in Cattle after Prostaglandin-Induced Luteolysis: Changes in Serum Hormones, Steroids in Follicular Fluid, and Gonadotropin Receptors. E Humbel, The amino acid sequence of human insulin-like growth factor I and its structural homology with proinsulin.J Biol Chem.
G Knight, C Glister, TGF-beta superfamily members and ovarian follicle development.Reproduction. G Knight, C Glister, Local roles of TGF-beta superfamily members in the control of ovarian follicle development.Anim Reprod Sci. Effects of various steroid milieus or physiological states on sexual behavior of Holstein cows. Timing of Insemination for Dairy Cows Identified in Estrus by a Radiotelemetric Estrus Detection System. W Alila, W Hansel, Origin of different cell types in the bovine corpus luteum as characterized by specific monoclonal antibodies.Biol Reprod.
Progesterone and conceptus elongation in cattle: a direct effect on the embryo or an indirect effect via the endometrium?
Effect of pregnancy and progesterone concentration on expression of genes encoding for transporters or secreted proteins in the bovine endometrium.
A Crowe, et alEffect of increasing progesterone concentration from Day 3 of pregnancy on subsequent embryo survival and development in beef heifers.Reprod Fertil Dev. E Spencer, O Sandra, E Wolf, Genes involved in conceptus-endometrial interactions in ruminants: insights from reductionism and thoughts on holistic approachesReproduction. It will grow well with a bit of lime in the soil, but is likely to become chlorotic if too much lime is present.
The Nissan GT-R features a newly developed Premium Midship package, including the world’s first independent transaxle 4WD developed independently by Nissan. It is a 191-amino acid, single chain polypeptide hormone which is synthesized, stored, and secreted by the somatotroph cells within the lateral wings of the anterior pituitary gland. In recent years, replacement therapies with human growth hormones (HGH) have become popular in the battle against aging. Examples of other causes of shortness often treated with GH are Turner syndrome, chronic renal failure, Prader-Willi syndrome, intrauterine growth retardation, and severe idiopathic short stature.
At the conclusion of the study all the men showed statistically significant increases in lean body mass and bone mineral, while the control group did not.
No other critical factors were affected, such as bone density, cholesterol levels, lipid measurements, maximal oxygen consumption, or any other factor that would indicate increased fitness Researchers also didn't discover any gain in muscle strength, which led them to believe that GH merely let the body store more water in the muscles rather than increase muscle growth. Identification of Potential Intrafollicular Factors Involved in Selection of Dominant Follicles in Heifers.Biol Reprod. E Echternkamp, The ovarian insulin and insulin-like growth factor system with an emphasis on domestic animals.Domest Anim Endocrinol. E Fortune, Proteolysis of Insulin-Like Growth Factor Binding Proteins-4 and-5 in Bovine Follicular Fluid: Implications for Ovarian Follicular Selection and Dominance.
C Wiltbank, Relationship between level of milk production and estrous behavior of lactating dairy cows.Anim Reprod Sci.
Progesterone-regulated changes in endometrial gene expression contribute to advanced conceptus development in cattle.Biol Reprod.
Its range extends in the Himalayas from Himachal Pradesh in India to southwest China and Burma.


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Carbon fiber, aluminum and steel components are assembled with a special high-precision process, which includes a series of vibration tests. Reported effects include decreased body fat, increased muscle mass, increased bone density, increased energy levels, improved skin tone and texture, and improved immune system function. GH can also be used in patients with short bowel syndrome to lessen the requirement for intravenous total parenteral nutrition. The authors of the study noted that these improvements were the opposite of the changes that would normally occur over a 10 to 20 year aging period. Some cows get steroids in their feed; others receive one or more hormones via a controlled-release implant in their ears. IntroductionGonadotropins are protein hormones secreted by the pituitary gland and include luteinizing hormone (LH) and follicle stimulating hormone (FSH). D Neill, Molecular cloning, sequencing, and characterizing the bovine receptor for gonadotropin releasing hormone (GNRH).Domest Anim Endocrinol. A Shupnik, Differential Gonadotropin-Releasing Hormone Stimulation of Rat Luteinizing Hormone Subunit Gene Transcription by Calcium Influx and Mitogen-Activated Protein Kinase-Signaling Pathways.Mol Endocrinol. A Beg, K Kot, Role of low circulating FSH concentrations in controlling the interval to emergence of the subsequent follicular wave in cattle.Reproduction.
E Mayo, Cellular Localization and Hormonal Regulation of Follicle-Stimulating Hormone and Luteinizing Hormone Receptor Messenger RNAs in the Rat Ovary.Mol Endocrinol.
T Armstrong, Estradiol-17beta biosynthesis in cultured granulosa cells from hypophysectomised immature rats; stimulation by follicle-stimulating hormone. Alterations in follicular IGFBP mRNA expression and follicular fluid IGFBP concentrations during the first follicle wave in beef heifersAnim Reprod Sci. E Spencer, et alSelect Nutrients and Their Associated Transporters Are Increased in the Ovine Uterus Following Early Progesterone Administration.Biol Reprod.
Cerasoides, like most members of this genus, is shallow rooted and is likely to produce suckers if the root is damaged. Despite the higher doses, side effects during treatment are rare, and vary little according to the condition being treated. Despite the fact the authors at no time claimed that GH had reversed the aging process itself, their results were misinterpreted as indicating GH was an effective anti-aging agent.
Association between surges of follicle-stimulating hormone and the emergence of follicular waves in heifers.J Reprod Fertil. F Roche, Follicular growth and serum follicle-stimulating hormone (FSH) responses to recombinant bovine FSH in GnRH-immunized anoestrous heifersAnim Sci. F Roche, Effects of Follicle-Stimulating Hormone With and Without Luteinizing Hormone on Serum Hormone Concentrations, Follicle Growth, and Intrafollicular Estradiol and Aromatase Activity in Gonadotropin-Releasing Hormone-Immunized Heifers.Biol Reprod. E Lamming, The detection and treatment of post insemination progesteroneinsufficieny in dairy cows. F Roche, P Lonergan, Evans ACO, et alThe repeatability of embryo survival, and the relationship between plasma progesterone in the early luteal phase and embryo survival in dairy heifers.
C Burghardt, M Palmarini, Pregnancy recognition and conceptus implantation in domestic ruminants: roles of progesterone, interferons and endogenous retrovirusesReprod Fertil Dev. G Wheeler, Endocrine changes associated with luteal regression in the ewe; the secretion of ovarian oestradiol, progesterone and androstenedione and uterine prostaglandin F2 alpha throughout the oestrous cycleJ Endocrinol. The pattern of secretion of LH pulses during an 8-h window early in the luteal phase (greater frequency, lesser amplitude), the mid-luteal phase (lesser frequency, lesser amplitude) and the follicular phase (high frequency, building to the surge) is indicated in the inserts in the top panel. J Smith, et alEffect of systemic progesterone concentration on the expression of progesterone-responsive genes in the bovine endometrium during the early luteal phase.Reprod Fertil Dev. The seed requires two to three months cold stratification and is best sown in a cold frame as early in winter as possible. Treatment increases animals' growth by 20%, so each cow in a feedlot typically gains 3 pounds (about Kg1.36) per day. F Ethier, Recruitment and development of the follicle; the roles of the transforming growth factor-? superfamily. Moreover, for each pound that it gains, it consumes 15% less feed than an untreated animal does. The scientists reported that hormone residues found in meat from these animals can disrupt the consumer's hormone balance, cause developmental problems, interfere with the reproductive system, and even lead to the development of cancer; (2) children and pregnant women are most susceptible to these negative health effects. The cycle consists of two discrete phases: the luteal phase (14–18 days) and the follicular phase (4–6 days). The luteal phase is the period following ovulation when the corpus luteum (CL) is formed (often further designated as met-estrus and diestrus), while the follicular phase is the period following the demise of the corpus luteum (luteolysis) until ovulation (often further designated as pro-oestrus and oestrus).
During the follicular phase, final maturation and ovulation of the ovulatory follicle occurs, the oocyte is released into the oviduct allowing the potential for fertilization.
The directive confirms the prohibition of substances having a hormonal action for growth promotion in farm animals. Gonadotropin regulation of follicle growth during the estrous cycleCattle are polyestrous animals and display estrous behavior approximately every 21 days. By October 2005 the Commission will present a report on the availability of alternative veterinary medicinal products; in April 2004, revelations that up to 90% of US veal calves are being fed synthetic testosterone illegally sent a shock wave through the meat industry, causing a government crackdown and new worries about the impact of hormones on the food supply.
In interviews with the media, veal industry officials said that calves have been fed growth hormones for decades.
These hormones function through a system of positive and negative feedback to govern the estrous cycle of cattle [1].
Officials with the Food and Drug Administration, however, say this has never been legal and the safety of this practice has not been tested. Its control of the estrous cycle is mediated via its actions on the anterior pituitary which regulates the secretion of the gonadotrophs, LH and FSH [4]. After transportation of GnRH from the hypothalamus to the pituitary gland via the hypophyseal portal blood system [5], GnRH binds to its G-protein coupled receptor on the cell surface of the gonadotroph cells [6].
This binding releases intracellular calcium which activates intermediaries in the mitogen activated protein kinases (MAPK) signaling pathway culminating in the release of FSH and LH from storage compartments in the cytoplasm [7]. FSH is only stored in secretory granules in the cytoplasm for short periods of time, whereas LH is stored for longer periods during the estrous cycle [8]. During the follicular phase of the estrous cycle there is a hormonal environment of basal progesterone due to the regression of the corpus luteum (CL). Only when serum progesterone concentrations are basal and LH pulse frequency increases to one per hour for 2–3 days does the DF ovulate [1]. Ovulation occurs 10–14 h after estrus and is followed by the luteal phase of the estrous cycle. The beginning of the luteal phase is also known as met-estrus and typically lasts 3–4 days. It is characterised by the formation of the CL from the collapsed ovulated follicle (corpus haemorragicum). During the di-estrous phase, progesterone concentrations remain elevated and recurrent waves of follicle development continue to be initiated by release of FSH from the anterior pituitary. However, these DFs that grow during the luteal phase of the estrous cycle do not ovulate, due to inadequate LH pulse frequency.The progesterone dominant luteal phase of the estrous cycle, through negative feedback, only allows the secretion of greater amplitude but less frequent LH pulses (one pulse per 3 to 4 hours) that are inadequate for ovulation of the DF [12].
Finally, during the pro-estrous period, progesterone concentrations decrease when the CL regresses in response to PGF secretion from the uterus [13].3. Gonadotropin regulation of final maturation of the pre-ovulatory follicle and ovulationThe growth, development and maturation of ovarian follicles are fundamental processes for high reproductive efficiency in farm animals. A fixed number of primordial follicles are established during fetal development with ovarian follicle growth taking a period of 3–4 months and categorized into gonadotropin independent and gonadotropin dependent stages [14].
Taken from [17].Each wave of growth involves emergence, selection and dominance followed by either atresia or ovulation of the DF.
Given that follicles are involved in the positive and negative feedback mechanisms of the hypothalamic–pituitary–gonadal (HPG) axis (estradiol and inhibins), these hormones have a governing role in the regulation of the estrous cycle of cattle.


The beginning of gonadotropin dependent follicle development is typified by the emergence of a follicle cohort typically consisting of 5–20 follicles ?5mm and is correlated with a transient increase in FSH concentrations [10, 18]. This marks the beginning of dependency of follicle growth on FSH [19] with FSH receptors (FSH-R) localized within the granulosa cells of the follicles by Day 3 of the follicle wave [20, 21]. This enables FSH to perform its required down stream signalling effects including promoting cellular growth and proliferation [22, 23]. These transient increases in FSH concentrations also leads to an increase in aromatase enzyme activity (P450arom; CYP19), in the granulosa cells of ovarian follicles, which converts androgen to estrogen [24].
As the DF is selected from the cohort of follicles, the diameter increases and it is recognized as the largest healthy follicle in the cohort [25]. This increase in size leads to an increase in follicular fluid estradiol and inhibin concentrations [24]. Dominance occurs when the the DF reaches 9 mm in diameter, and it actively suppresses FSH, thus preventing further follicle wave emergence until the DF either undergoes atresia or ovulated. The increase in estradiol concentrations in concert with inhibin are the key endocrine signals that suppress FSH concentrations from the anterior pituitary gland via negative feedback reducing FSH to basal concentrations [10, 26, 27]. The selected DF becomes increasingly responsive to LH [27] and continues growth in the face of decreasing FSH concentrations. Irrespective of the stage of the estrous cycle during which follicles develop, the switch from FSH [18] to LH dependency [28] is propagated through the presence of LH receptors (LH-R) on the granulosa cells [29].
LH-R are localised to the theca and granulosa cells of healthy follicles, at different stages of follicle development [20]. As the follicle grows, the theca cell LH-R increases and LH-R is acquired by the granulosa cells of the follicle undergoing selection to become the DF [29-31]. Moreover, evidence suggests transient increases in circulating LH concentrations that occur at or around the time of follicle selection [32], allows the DF to continue E2 production and grow in the face of declining FSH concentrations [33].
The production of high concentrations of estradiol is a defining characteristic of the DF [33, 34] and prior to visible differences in follicle diameter; the putative DF has greater follicular fluid concentrations of estradiol compared with other follicles in its cohort [10, 35, 36].
Production of estradiol from growing follicles is dependent on sufficient LH pulse frequency [38, 39].
The binding of LH to its receptors in the theca cells drives the conversion of cholesterol to testosterone through a series of catalytic reactions [40]. Testosterone, once produced in the theca cells, diffuses out into the granulosa cells where it is converted to estrogens by the aromatase enzyme [40]. Estradiol not only has a local effect on follicle development, but it also has a systemic role via a positive feedback mechanism to the hypothalamus and pituitary gland.
During the follicular phase of the estrous cycle, when progesterone concentrations are basal, this large concentration of estradiol produced by the pre-ovulatory DF induces a GnRH surge from the hypothalamus. The resulting LH surge is of sufficient amplitude and frequency to stimulate final maturation and ovulation of the DF [10]. The increased estradiol concentrations also induces expression of estrous behavour, required for successful mating [41]. Other intra-ovarian produced factors play a role in regulating the estrous cycle either indirectly by altering the synthesis of estradiol or via direct negative feedback mechanisms to the hypothalamus and the anterior pituitary gland.
The insulin like growth factor (IGF) super-family consisting of its two ligands IGF-I and IGF-II [42-44], two receptors IGFR-I and IGFR-II [45], and it numerous binding proteins and proteases (IGFBP 1-6, pregnancy-associated plasma protein-A: PAPP-A) are responsible for the bioavailability of IGF-1 in the ovarian follicle. The bioavailability of IGF-I contributes to the growth, proliferation and steroidogenic capacity of the future DF [36, 46, 47], indirectly affecting the estradiol induced negative feedback mechanism to the hypothalamus and pituitary.
This in addition to early acquisition of LH receptors by the granulosa cell layer of the follicle undergoing selection are considered to be the main mechanisms facilitating the process of follicle selection [48]. The ligand members of this super-family were first identified in follicular fluid through their modulation of secreted FSH [49].
Activin can increase the production of estradiol in follicular fluid [50] whereas follistatin impedes activins’ positive steroidogenic effects, both of which can alter the estradiol feedback mechanism to the hypothalamus and pituitary [51]. Inhibins which have been detected in granulosa cells in cattle play a role in the suppression of FSH secreted in the anterior pituitary also regulating the oestrous cycle [52].4. Standing to be mounted by a bull or herd mate is the primary and most definitive sign of oestrus in cattle. Estrogen, specifically, estradiol, is the primary signal to the brain that induces expression of estrus, but only in the absence of progesterone [54]. It appears that stressors which elevate blood concentrations of cortisol are capable of delaying or blocking the pre-ovulatory LH surge and affecting the expression of estrus without altering pro-oestrous concentrations of blood oestradiol (see review by [55]).
For heifers it would appear that the duration of standing estrus is somewhat longer, 12–14 h [56]. For beef cows, kept indoors, the average duration of standing estrus has been reported to be less than 8.5 h [56].
Both the duration of standing estrus and intensity of estrous expression are affected by a range of environmental factors including under foot surface type, size of the sexually active group and the presence of a bull [56]. Breaks or quiescent interludes in standing activity have also been observed in 30% of dairy cows at [58] while breaks with an average duration of 2.6 h in 67% of beef heifers have been recorded [59]. There is no evidence from dairy cows [60], beef cows or heifers [56] that either the onset of standing estrus or end of estrus follows any distinct diurnal pattern.5.
Gonadotropin regulation of Corpus luteum function The CL originates from the cells of the ovulatory follicle. LH, the major luteotropic hormone in cattle [61], is responsible for stimulating luteinization of the theca and granulosa cells of the pre-ovulatory follicle into luteal cells [62]. The function of the CL is to produce sufficient concentrations of progesterone throughout the luteal phase of the estrous cycle to maintain pregnancy (if a conceptus is present) and during pregnancy, to decrease gonadotropin secretion and prevent behavioral oestrus occurring. Progesterone is required for the maintenance of pregnancy with many studies reporting a positive association between progesterone concentrations and the probability of embryo survival [63-66].
The proposed mechanisms by which progesterone affects embryo survival are indirect, not acting on the embryo itself but via effects on the uterine endometrium [67, 68]. Available evidence in both cattle and sheep, has identified that sustained increased concentrations of progesterone during the luteal phase of the estrous cycle alters the expression pattern of genes in the uterus [69-73] which in turn alters the composition of the uterine histotroph i.e. Moreover, alterations in systemic progesterone during the early luteal phase have been shown to have significant effects on conceptus elongation [67, 71, 74].
During the mid-luteal phase, these sustained high concentrations of circulating progesterone down regulate the nuclear progesterone receptor in the luminal epithelium of the endometrium [75].
This is a critical switch in allowing the synchronous increase or decrease in genes of the endometrium that are required to initiate uterine receptivity – regardless of the pregnancy status of the animal [76]. If, by Day 16 of the estrous cycle, the maternal recognition of pregnancy signal (interferon tau) has not been detected in sufficient quantities, luteolysis of the CL occurs. PGF is secreted by the uterus in the bovine [77] and is the major luteolytic hormone in ruminants [78-80]. Oxytocin receptors in the uterus binds oxytocin which propagates the episodic secretion of PGF from the uterus. PGF then mediates the luteolytic mechanism via countercurrent exchange between the uterine vein and the ovarian artery (Fig. This reduces circulating progesterone concentrations, estradiol concentrations increase and GnRH in the hypothalamus is stimulated as the animal enters the follicular phase of the estrous cycle.6. ConclusionsThe estrous cycle in cattle is typically 18–24 days in duration, with estrous behavior expressed for a 2–24-h period during the late follicular phase. During normal estrous cycles there are typically two to three and occasionally four waves of follicular growth each involving a period of emergence, selection and dominance followed by either atresia or ovulation of the DF. The gonadotropin hormones FSH and LH are the main regulators of folliculogenesis and steroidogenesis with LH being the major luteotrophic hormone.



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