Relationship between growth hormone (GH) pulses in the peripheral circulation and GH-releasing hormone and somatostatin profiles in the cerebrospinal fluid of goats

Growth hormone (GH) is secreted in a pulsatile manner, but the underlying mechanisms of GH pulse generation remain to be resolved. In the present study, we investigated the relationship between GH pulses in the peripheral circulation and GH-releasing hormone (GHRH) and somatostatin (SRIF) profiles in the cerebrospinal fluid (CSF) of male goats. The effects of an intracerebroventricular (icv) injection of neuropeptide Y (NPY), galanin and ghrelin were also analyzed. Blood and CSF samples were collected every 15 min for 8 hr from the jugular vein and third ventricle, respectively. GH pulsatility in the goat was found to consist of distinct large pulses of 5 hr periodicity and small pulses of 1 hr periodicity. GHRH and SRIF in the CSF fluctuated in a pulsatile manner with 1 hr periodicity, and most of the descending phase of SRIF pulses were associated with the initiation of GH pulses. Icv injections of NPY, galanin and ghrelin stimulated GHRH release without affecting SRIF release. In addition, NPY suppressed, and galanin and ghrelin induced large GH pulses, although ghrelin was much more effective than galanin. These results suggest that an hourly fall in SRIF is involved in generating intrinsic circhoral rhythm of GH pulsatility. The mechanisms underlying the generation of large GH pulses of 5 hr periodicity remain unknown, while direct action of NPY and/or ghrelin on the pituitary might be involved.     

Temporal patterns of growth hormone in blood and seminal plasma of mature dairy bulls

This study was conducted to determine if growth hormone (GH) concentration in bovine seminal plasma would be proportional to but less variable than blood plasma GH. The relationship between GH in blood and seminal plasma was also examined critically. Blood samples were collected at 15-min intervals for 5.75 h, while semen was collected at 30-min intervals over the same time period. Average seminal plasma GH concentrations were 3.2 times higher (P less than .05) than blood plasma GH concentrations (40.4 +/- 15.8 ng/ml vs 12.6 +/- 1.2 ng/ml, respectively). The within animal correlation between blood and seminal plasma was consistently low and nonsignificant (P greater than .05). Overall blood plasma GH and seminal plasma GH concentrations were weakly correlated (r = .418; P greater than .05) among bulls. A predictable relationship between blood and seminal plasma GH concentration does not exist under the conditions of this study.     

Suppressing effects of short-chain fatty acids on growth hormone (GH)-releasing hormone-induced GH release in isolated anterior pituitary cells of goats.

The involvement of tetrodotoxin-sensitive Na+ channels and receptor-operated nonspecific Ca2+ channels, and the effects of short-chain fatty acids, on growth hormone (GH) release induced by GH-releasing hormone (GHRH) were investigated in cultured and freshly isolated caprine anterior pituitary cells. In 3-d cultured cells in Dulbecco's modified Eagle's medium, an increase in GH release induced by GHRH (10 nmol/l) was moderately, but significantly, reduced by a voltage-sensitive Na+ channel antagonist tetrodotoxin (1 micromol). The GHRH-induced GH increase, which was not affected by a simultaneous addition of a receptor-operated nonspecific Ca2+ channel antagonist tetramethrine (0.1 mmol/l), was significantly reduced by a voltage-sensitive L-type Ca2+ channel antagonist nifedipine (1 micromol/l). Propionate and butyrate at 10 mmol/l, however, not only suppressed basal GH release but also significantly reduced the GH increase induced by 10 nmol/l of GHRH. The inhibitory action of these acids was also reproduced by an addition of beta-hydroxy butyrate (10 mmol/l) and octanoate (10 mmol/l). In freshly isolated and perifused cells, butyrate (10 mmol/l) as well as somatostatin (100 nmol/l) significantly reduced the GH increase induced by GHRH. From these findings we conclude that tetrodotoxin-sensitive Na+ channels and voltage-dependent L-type Ca2+ channels are involved in the cellular mechanism for GHRH-induced GH release, and that short-chain fatty acids such as propionate and butyrate have a direct action on somatotrophs to reduce basal and GHRH-induced GH release, in caprine somatotrophs.     

Ovarian follicle populations and ovulation rates of Finnish Landrace cross ewes in different nutritional states and associated profiles of gonadotrophins, inhibin, growth hormone(GH) and insulin-like growth factor-I

Effects of ewe body condition and level of feed intake on ovarian follicle populations and ovulation rates were studied in three groups of 12 to 15 Finnish Landrace cross ewes. Ewes were fed to achieve target body condition scores of either 2.0 (low, LM) or 2.5 (moderate, MM) and then fed a complete pelleted ration for three weeks before study so that they maintained liveweight. Ewes of a third group were also fed to achieve condition scores of 2.5 and then fed ad libitum (MAL). Neither ewe body condition nor level of feed intake significantly affected either the number of large follicles (greater than or equal to 4 mm diameter) or the ovulation rate. The proportion of estrogenic follicles was lower in MM than LM ewes (0.77 vs. 0.96; P less than 0.05), but there was no difference in this proportion between MAL and MM ewes. Insulin-like growth factor-I (IGF-I) concentrations in the follicular fluid were unaffected by ewe body condition or level of feed intake despite significant treatment differences in circulating concentrations. Inhibin concentrations were higher in estrogenic follicles of LM compared with MM ewes, and this difference was reflected in circulating profiles. Treatment differences in LH profiles were not associated with any difference in follicle populations or ovulation rate. There were no consistent treatment effects on FSH concentrations. It is concluded that the roles of inhibin and IGF-I in the control of follicle development cannot be adequately assessed on the basis of circulating concentrations alone and that there is a need to measure intrafollicular hormone profiles and associated effects on follicle physiology.     

Changes in plasma non-esterified fatty acids, glucose and alpha-amino nitrogen and their relationship with body weight and plasma growth hormone in growing buffaloes (Bubalus bubalis)

A study was undertaken to investigate the changes of plasma non-esterified fatty acids (NEFA), glucose and alpha-amino nitrogen and their relationship with age, body weight (BW) and plasma growth hormone (GH) in growing buffaloes. For the purpose, six growing female Murrah buffalo calves of 6-8 months of age were selected on the basis of their BW and fed according to Kearl standard (Nutrient Requirements of Ruminants in Developing Countries, International Feedstuffs Institute, Utah State University, Utah, USA, 1982, p. 89) for growing buffaloes (target growth rate 500 g/day) to meet energy and protein requirement of the animals. Blood samples collected at fortnight intervals for 1 year were analysed for plasma NEFA, glucose, alpha-amino nitrogen and GH. The animals were also weighed at fortnight intervals. Plasma NEFA and glucose levels were found to decrease (p < 0.01) with age. Unlike plasma NEFA and glucose, plasma alpha-amino nitrogen level increased (p < 0.01) as the buffaloes become older. Plasma NEFA and glucose concentrations in growing buffaloes were found to be positively correlated with plasma GH (r = 0.379 and 0.420 respectively), but these were non-significant (p > 0.01). However, plasma NEFA and glucose showed a good correlation (p < 0.01; r = 0.780 and 0.652 respectively) with plasma GH per 100 kg live weight. Plasma alpha-amino nitrogen exhibited non-significant (p > 0.01) negative correlation (r = -0.295) with plasma GH but a negative correlation (p < 0.01; r = -0.641) with GH per 100 kg BW. So, plasma metabolites showed a definite pattern of change during growth and these have a significant (p < 0.01) correlation with plasma GH per 100 kg BW than GH.     

Progestin-induced growth hormone (GH) production in the treatment of dogs with congenital GH deficiency

The recent demonstration of the ability of progestins to induce the expression of the growth hormone (GH) gene in the mammary gland of dogs and cats opens possibilities for the treatment of some forms of GH deficiency with progestins. Therefore, one male and one female German shepherd dog with congenital dwarfism because of a pituitary anomaly were treated with subcutaneous injections of medroxyprogesterone acetate (MPA) in doses of 2.5–5.0 mg per kg body weight, initially at 3-wk intervals and subsequently at 6-wk intervals. In both dogs, body sizes increased and a complete adult hair coat developed. Undesirable side-effects were recurrent periods of pruritic pyoderma in both dogs and cystic endometrial hyperplasia with mucometra in the female dog. Parallel with the physical improvements, plasma insulin-like growth factor I concentrations rose sharply. Plasma GH concentrations tended to rise, but never exceeded the upper limit of the reference range. Nevertheless, one of the dogs developed slight acromegalic features, possibly because mammary GH, unlike pituitary GH, is released evenly throughout the day. Even moderate increases in circulating GH concentration may, therefore, give rise to overexposure. It is concluded that long-term treatment with MPA can be used as an alternative for heterologous GH in the treatment of congenital GH deficiency in the dog.     

Functional expression of the free fatty acids receptor-1 and -4 (FFA1/GPR40 and FFA4/GPR120) in bovine endometrial cells

Veterinary Research Communications - Endometrial epithelial cells play a key defensive role as part of the innate immune response of cow uterus. An association between risk of acquiring infectious...     

Relationship between testicular measurements, body weight and semen quality in young dairy bulls.

Dairy bulls, 322 Ayrshires (Ay) and 85 Friesians (Fr), were studied at the age of 11 months. Of the bulls, 286 Ay-bulls and 80 Fr-bulls produced semen of acceptable quality for use in A.I. Scrotal circumference, tonometer measure, scrotal fold thickness, 1-year body weight and testicular palpation were used to predict unsuitable bulls for A.I. Non-return rate was used as a measure of fertility. Scrotal fold thickness and 1-year weight had no significant correlation with fertility or semen quality. Scrotal circumference had a significant positive correlation with fertility. Tonometer ratio had a significant negative correlation with fertility. Testicular palpation was the best basis for predicting bulls with poor semen quality in this study. Twelve bulls were recorded as having testicles of different sizes, 1 testicle being more than 20% bigger than the other. Only 2 of these 12 bulls produced semen of acceptable quality. One of these 2 bulls was, after slaughter, diagnosed as having a hereditary testicle disease. Friesians were shown to have significantly higher fertility than Ayshires.     

Relationship between growth and plasma concentrations of ghrelin and growth hormone in juvenile beagle dogs

Although the release of growth hormone (GH) is known to be regulated mainly by GH-releasing hormone (GHRH) and somatostatin (SRIF) secreted from the hypothalamus, ghrelin also may be involved in GH release during juvenile period. We have examined plasma concentrations of acylated ghrelin, desacyl ghrelin, and GH in juvenile beagle dogs. Plasma acylated and desacyl ghrelin levels changed through aging; however, there was no closely correlation between ghrelin, body weight and circulating GH levels during juvenile period. The increase in body weight was essentially linear until 8 months of age, whereas plasma GH concentrations exhibited bimodal peaks for the meanwhile. The results suggest that ghrelin may not play internal cueing in GH secretion in juvenile beagle dogs.     

Peripheral T lymphocyte changes in neonatal piglets: Relationship with growth hormone (GH), prolactin (PRL) and cortisol changes

Taking into account the role played by the neuroendocrine network in affecting the early development of the immune response, the present study aims to assess neonatal immunity in piglets by testing peripheral lymphocyte age-related changes in relationship to plasma levels of some relevant immunoregulatory hormones, such as growth hormone (GH), prolactin (PRL) and cortisol. For this purpose, we studied the peripheral lymphocyte age-related changes in relationship to plasma levels of GH, PRL and cortisol in conventional piglets from birth (day 0) to 41 days of age. A significant decrease was observed in the total number of lymphocytes at day 0, with a subsequent constant increment up to 41 days of age. Concomitantly, the number of T cell subsets (mainly CD8(+) cells and double positive CD4(+)CD8(+)) was low at birth, with strong increments between the 19th and 41st days of life. The CD4(+) T cell number subset was less diminished at birth than that of CD8(+), albeit with significant increments in the post-weaning period. Of interest, gammadelta T cells, which are more involved in innate immune efficiency, displayed the same trend as CD8(+) T cells from birth to the 41st day of life. From day 0 up to the 19th day, significant inverse correlations were found between T cell subsets and GH or PRL or cortisol, albeit with more significant inverse correlations with cortisol. The high levels of GH and PRL in the pre-weaning period may be due to the fact that they have to counteract the cortisol-mediated negative effect on lymphocyte production and development. These findings suggest that stress condition occurs at birth with decreases in the immune parameters, in the same way as in human newborns, with a subsequent gradual normalisation and immune development, as shown by decreased cortisol, GH and PRL normalisation and concomitant increments in T cell subsets.     

Relationship of thyroid status to growth hormone and insulin-like growth factor-I (IGF-I) in plasma and IGF-I mRNA in liver and skeletal muscle of cattle

Steers were made hyperthyroid or hypothyroid to study the effects of physiological alterations in thyroid hormone status on plasma growth hormone (GH) profiles, plasma insulin-like growth factor-I (IGF-I) concentrations, and relative abundance of IGF-I mRNA in skeletal muscle and liver. Eighteen yearling crossbred steers (360 to 420 kg) were randomly allotted to hyperthyroid (subcutaneous injection 0.6 μg/kg BW L-thyroxine for 10 d), hypothyroid (oral thiouracil; 0.25% diet plus 12.5 g capsule/d for 17 d), or control (subcutaneous injection 0.9% NaCl) treatment groups. Blood samples were taken for measurement of GH, IGF-I, thyroxine (T₄) and triiodothyronine (T₃) by RIA. Samples of liver and skeletal muscle were taken by biopsy for measurement of IGF-I mRNA by solution hybridization. Steers receiving thiouracil had 57 and 53% (P<.05) lower T₄ and T₃, respectively, than control steers (84.1 and 1.7 ng/ml). The hyperthyroid steers had 228 and 65% greater (P<.05) T₄ and T₃ than control steers. Neither increased nor decreased thyroid status had any significant effects on plasma GH profiles, liver IGF-I mRNA, or plasma concentration of IGF-I. There was no effect of thyroid hormone alteration on skeletal muscle IGF-I mRNA concentrations. The results of this study suggest that short-term changes in thyroid status of cattle had no major impact on the GH-IGF-I axis or skeletal muscle IGF-I mRNA.     

Inability of heterologous growth hormone (GH) to regulate GH binding protein in GH-transgenic swine

Transgenic pigs expressing bovine, ovine, or human growth hormone (GH) structural genes fused to mouse metallothionein-I (mMT-bGH), ovine MT (oMT-oGH), or mouse transferrin (mTf-hGH) promoters were used to study the effects of GH on the regulation of serum GH-binding protein (GHBP). In the 14 transgenic pigs studied, circulating concentrations of heterologous GH ranged from 15 to 2,750 ng/mL. Using chromatographic methods, specific binding of GH was detected in serum from normal pigs but was undetectable in serum from all the transgenic pigs used, probably as a result of the high serum concentrations of heterologous GH present in these animals. Thus, to avoid interference of binding by high GH concentrations, serum samples were subjected to immunoblotting using a specific anti-GHBP antibody. A specific 54-kDa band was detected in normal pig serum as well as in sera from mMT-bGH, oMT-oGH, and mTf-hGH pigs. Additionally, sera from transgenic mMT-bGH pigs and their sibling controls were subjected to immunoprecipitation with an anti-GHBP antibody followed by immunoblotting with the same antibody. With this technique, we detected two specific bands of 53 and 45 kDa that could represent different degrees of glycosylation of GHBP. As determined by densitometric analysis the amount of GHBP in transgenic pig sera was similar to that detected in sera of the respective control animals. The amount of circulating GHBP remained unchanged even in oMT-oGH and mTf-hGH pigs that were exposed from birth to circulating concentrations of GH as high as 2,750 ng/mL. Thus, we conclude that heterologous GH do not act as modulators ofthe serum GHBP in pigs.     

Effect of triglycerides on growth hormone (GH)-releasing factor-mediated GH secretion in newborn calves

The effect of triglycerides (Tg) on GRF-mediated GH secretion was examined in 2 groups of twelve ten-day old male calves. Twelve calves were intravenously infused with a lipid-heparin solution (5 mg Tg and 0.3 IU heparin/kg body wt/min for 90 min). The twelve control calves received in the same way, the same volume of saline. Thirty minutes after the start of infusion, GRF 1–29 (human amide, 0.16 μg/kg body wt) was intravenously injected in six animals of each group.

Mean plasma GH levels reached peak concentrations in the 2 groups 5 min after GRF injection. However the area under the GH response curve, when lipid-heparin was given, was significantly diminished compared to the response when saline was given. In the same time, lipid-heparin treatment increased plasma SRIF concentration. These data suggest that an increase in plasma Tg concentration, induced by lipid-heparin infusion, inhibits GRF-mediated GH secretion, possibly through stimulation of SRIF secretion.     

Response of plasma fatty acid profiles to changes in dietary n‐3 fatty acids and its correlation with erythrocyte fatty acid profiles in dogs

An elevated level of long‐chain n‐3 fatty acids (FA) in tissue membranes has a positive influence on the progression and treatment of many diseases. Therefore, dietary supplementation of n‐3 FA is recommended in some diseases. Even though n‐3 FA are absorbed readily from the diet, their incorporation into tissues may be compromised in diseased animals. In a clinical setting, it is desirable to monitor the success of dietary intervention. Plasma FA as well as erythrocyte membrane (EM) FA can be used to monitor dietary FA intake. This study compares FA from EM and plasma with regard to their reaction time and reliability for monitoring dietary changes of tissue FA profiles in dogs. Thirty dogs were divided into three groups and fed for 12 weeks. The control group (CONT) was fed a commercial standard diet low in n‐3 FA. One group received the standard diet and 85 mg/kg body weight of a docosahexaenoic acid (DHA) concentrate (ADD). The third group was fed a commercial dog food containing fish oil (FO), which is rich in eicosapentaenoic acid (EPA). EM and plasma FA profiles were analysed by GC separately. Data on EM FA were published recently. n‐3 FA in plasma reached the new level after 2 weeks (8 weeks in EM). Dietary differences between DHA and EPA are obvious after 1 week already. The concomitant decrease in plasma n‐6 FA differed between ADD and FO. In general, the correlation of n‐6 FA between plasma and EM was low. We therefore conclude that analysis of plasma FA is sufficient for monitoring a diet‐induced increase in tissue n‐3 FA in dogs. However, EM FA should be analysed if the effect of dietary intervention on tissue n‐6 FA is important.     

Relations between plasma acetate, 3-hydroxybutyrate, FFA, glucose levels and energy nutrition in lactating dairy cows.

To clarify the implication of an energy nutrition on a metabolic alteration with advancing lactation, total 270 blood samples were taken from 16 lactating dairy cows. Amounts of dietary allowance and the refusals were measured daily, and the energy (TDN) intakes and a satisfaction (energy balance) of each cow were estimated. Plasma acetate, 3-hydroxybutyrate (3-HB), free fatty acid (FFA) and glucose levels were estimated. The data were divided into 3 groups depending on the days in milk; early (up to 70 days postpartum), mid (71 to 140 days), and late (after 141 days) lactation. There were many cases of higher FFA level in early lactation, especially with declining acetate and glucose levels. There were proportional elevations of 3-HB in connection with FFA levels in many samples of early lactation, though the 3-HB increased independently of FFA levels in the most cases of the mid and late lactations. Plasma 3-HB levels increased in many cases of decreased glucose level, especially in the early lactation. Plasma acetate level correlated positively with 3-HB level, but not correlated with glucose level. Higher FFA level and elevation of FFA/3-HB ratio were observed in the conditions of negative energy balance. This implies the metabolic importance of FFA in a ketogenesis of the early lactation.     

Calcium salts of fatty acids with varying fatty acid profiles in diets of feedlot-finished Bos indicus bulls: impacts on intake,digestibility, performance,and carcass and meat characteristics

We hypothesized that the inclusion of calcium salts of fatty acid (CSFA) into the diets and the fatty acid (FA) profile of the supplements would impact performance and meat characteristics of Bos indicus bulls. Hence, the objective was to evaluate the effects of CSFA profiles on intake, body weight (BW), carcass, and meat characteristics of feedlot-finished B indicus bulls. Fifty-three Nellore bulls [initial BW 315 ± 5.9 kg and 20 ± 2 mo] were used. At the beginning, 6 bulls were randomly chosen and slaughtered for determination of their BW composition, and the remaining 47 bulls were evaluated during a 140-d experimental period. The bulls were placed in individual pens, blocked according to initial BW and randomly allocated to 1 of the 3 following treatments: (1) control diet containing sugarcane bagasse, ground corn, citrus pulp, peanut meal, and mineral–vitamin mix (CON), (2) CON with the addition of 3.3% of CSFA from soybean oil (CSO), or (3) CON with the addition of a mixture of 3.3% of CSFA from palm, soybean, and cottonseed oils (CPSCO). Diets were offered ad libitum and formulated to be isonitrogenous. Bulls supplemented with CSFA had a greater (P < 0.01) final BW, dry matter intake, average daily gain (ADG), feed efficiency (FE), and FA intake vs. CON. Among carcass parameters, CSFA-supplemented bulls had greater (P < 0.01) carcass ether extract concentration vs. CON bulls. When the CSFA profile was evaluated (CSO vs. CPSCO), CPSCO bulls had a better (P ≤ 0.03) FE, carcass ADG, and hot carcass weight (HCW) vs. CSO bulls. The FA intakes differed among CSFA treatments, as the total saturated, palmitic, and oleic FA intakes were greater for CPSCO (P < 0.01), whereas lower intakes of total unsaturated and polyunsaturated FA (P < 0.01) were observed for CPSCO vs. CSO. Samples from the Longissimus muscle contained greater palmitoleic (P = 0.01) and reduced linoleic (P = 0.02) FA concentrations in CSFA-supplemented bulls vs. CON bulls. In agreement with the FA intakes, CPSCO-supplemented bulls had a greater (P ≤ 0.05) unsaturated FA concentration vs. CSO in Longissimus muscle. In summary, CSFA supplementation improved the performance of finishing B. indicus bulls vs. CON. Moreover, the inclusion of CSFA from palm, soybean, and cottonseed oil benefited the FE, carcass ADG, and HCW compared with the inclusion of CSFA from soybean oil, demonstrating the potential of specific FA for improving the performance and meat quality of B. indicus bulls.     

Oral administration of peptidergic growth hormone (GH) secretagogue KP102 stimulates GH release in goats

To assess the oral activity of KP102 (also known GHRP-2) on growth hormone (GH) release in ruminant animals, 5 or 10 mg/kg body weight (BW) of KP102 dissolved in saline was orally administered twice at 2 hr-intervals to either 1- or 3-mo-old goats (n = 5-6). Plasma GH concentrations in the 1-mo-old goats were elevated at 15 min after the first administration of both 5 and 10 mg/kg BW of KP102. Significant elevation of GH concentrations continued until 180 min after 10 mg/kg BW of KP102, whereas the elevated GH levels after the administrations of 5 mg/kg BW of KP102 subsided to basal concentrations within 90 min. The second administration of 10 mg/kg BW of KP102 failed to elevate the GH concentration, but 5 mg/kg BW of KP102 abruptly stimulated GH release. Plasma GH concentrations in the 3-mo-old goats were also significantly elevated after the administration of both 5 and 10 mg/kg BW of KP102. The plasma GH responses to 5 and 10 mg/kg BW of KP102 were almost identical. The elevated GH levels after the first administration of KP102 tended to be maintained throughout the experiment, and a transient increase in plasma GH levels was observed after the second administration. However, the stimulatory effect of KP102 on GH release in the 3-mo-old goats was small and less abrupt than that in the 1-mo-old goats. The concentrations of insulin-like growth factor-I were not increased by KP102 during the brief sampling periods used in this experiment. These results show that the oral administration of the peptidergic GH secretogogue KP102 stimulates GH release in a ruminant species, and that the oral activity of KP102 on GH release is modified by the age.     

Relationship of growth hormone, prolactin and thyrotropin secretion to individual and progeny performance of Hereford bulls

Six Hereford bulls from a line selected with an index of body weight and muscling score were compared with six bulls from a control line to determine if increased growth in the selected line was associated with changes in plasma levels of growth hormone (GH), prolactin (PRL) and (or) thyrotropin (TSH). The predictive value of sire hormone data for growth rate of progeny was also evaluated. Bulls of the index line were heavier at birth (P less than .02) and had higher postweaning daily gains (P less than .01) than bulls of the control line. Blood samples were collected from bulls (ages 2 and 3 yr) at 15-min intervals for 8 h. Overall plasma GH concentrations were higher (P less than .03) in the index bulls than those in the control line. All characteristics of PRL secretion examined tended to be higher in the index bulls, but only mean overall and baseline differences approached significance (P less than .10). There were no significant differences in measures of TSH secretion between lines. Sire differences in hormone characteristics accounted for significant amounts of variation in birth weight and postweaning gain of progeny, but not in gain to weaning. Also, when considered jointly, hormone characteristics generally added significantly to predictions that used sire growth rate alone. The results suggest that serum hormone characteristics in parents may provide additional predictors of growth rates of progeny.