Ovulation and ovulation rate in ewes under grazing conditions: factors affecting the response to short-term supplementation.

The relationships between ovulation rate and nutrition remain confused, probably because of uncontrolled variation in experimental conditions. To help resolve the problem, we analyzed data from 20 experiments conducted between 2002 and 2016, in Uruguay with grazing ewes. All experiments were carried out by a single laboratory under comparable conditions of experimental design and measured variables. The studies used a total of 3 720 ewes, of purebred Corriedale, Polwarth, or East Friesian x Polwarth genotypes. In all experiments, a control group grazed native pastures and extra nutrition was provided to the treatment groups using either improved pastures or supplements. Ovulation rate was measured by counting corpora lutea using laparoscopy or rectal ultrasound or by counting foetuses at ultrasound on day 45 of gestation. For statistical analysis, data were grouped according to nutritional treatment (control or supplemented) and, within these groups, type of supplement to provide energy or protein (protected or not from rumen degradation). Across all experiments, 92-99% of the ewes ovulated and the effects of diet, length of supplementation, and initial live weight and genotype are reported. Within diets, ovulation was most affected by overall energy intake during supplementation (P < 0.01). Ewes that grazed native pastures supplemented with protein supplements had higher ovulation rates (P < 0.05) than control ewes grazing only native pastures. The addition of tannins to the protein supplement, to protect it from degradation in the rumen, did not further increase the ovulation rate. In unsupplemented ewes that had access to legume pastures, ovulation rates did not increase when the legume pasture was rich in tannins although only ewes that grazed tanniniferous legumes had marginally higher ovulation rates than the control ewes (P < 0.05). When ewes grazing native pastures were supplemented with energy, their ovulation rate did not increase above those of nonsupplemented ewes. Live weight at the start of supplementation also affected ovulation rate. We conclude that ovulation was most affected by overall energy intake, whereas the factors that affected ovulation rate during short-tern nutritional supplementation were intake of protein from highly digested supplements or dietary protein protected from ruminal degradation.

Sheep deficient in vitamin E preferentially select for a feed with a higher concentration of vitamin E.

Given the capacity of ruminants to modify diet selection based on metabolic needs, we hypothesised that, when given a choice, lambs experiencing a vitamin E deficiency would consume more of a vitamin E-enriched feed than lambs not deficient in vitamin E. Fifty-six Dohne Merino lambs were divided into two groups and fed either a vitamin E-deficient diet over 40 days to induce low plasma vitamin E or a vitamin E-enriched diet to induce high plasma vitamin E. The lambs were then offered a choice of vitamin E-enriched and vitamin E-deficient pellets. For half of the animals, the enriched diet was paired with strawberry flavour and the deficient diet was paired with orange flavour, while the reverse pairings were offered to the others. Lamb preference for the diets was measured daily for the following 15 days. There was a three-way interaction between the high and low vitamin E treatment groups×vitamin E content and type of flavour in the feed×time (days). The lambs preferred pellets flavoured with strawberry but this preference changed to orange flavour in vitamin E-deficient lambs if the orange flavour was paired with high vitamin E. Lambs without a deficiency continued to prefer strawberry-flavoured pellets, regardless of the vitamin E concentrations in the pellets. It is possible that self-learning contributed to the low vitamin E group of lambs changing preference to orange flavour in order to consume more vitamin E, presumably to remediate the deficiency.

Colostrum production in ewes: a review of regulation mechanisms and of energy supply.

In sheep production systems based on extensive grazing, neonatal mortality often reaches 15% to 20% of lambs born, and the mortality rate can be doubled in the case of multiple births. An important contributing factor is the nutrition of the mother because it affects the amount of colostrum available at birth. Ewes carrying multiple lambs have higher energy requirements than ewes carrying a single lamb and this problem is compounded by limitations to voluntary feed intake as the gravid uterus compresses the rumen. This combination of factors means that the nutritional requirements of the ewe carrying multiple lambs can rarely be met by the supply of pasture alone. This problem can overcome by supplementation with energy during the last week of pregnancy, a treatment that increases colostrum production and also reduces colostrum viscosity, making it easier for the neonatal lamb to suck. In addition, litter size and nutrition both accelerate the decline in concentration of circulating progesterone that, in turn, triggers the onsets of both birth and lactogenesis, and thus ensures the synchrony of these two events. Furthermore, the presence of colostrum in the gut of the lamb increases its ability to recognize its mother, and thus improves mother-young bonding. Most cereal grains that are rich in energy in the form of starch, when used as supplements in late pregnancy will increase colostrum production by 90% to 185% above control (unsupplemented) values. Variation among types of cereal grain in the response they induce may be due to differences in the amount of starch digested post-ruminally. As a percentage of grain dry matter intake, the amount of starch entering the lower digestive tract is 14% for maize, 8.5% for barley and 2% for oats. Supplements of high quality protein from legumes and oleiferous seeds can also increase colostrum production but they are less effective than cereal grains. In conclusion, short-term supplementation before parturition, particularly with energy-rich concentrates, can improve colostrum production, help meet the energy and immunological requirements for new-born lambs, and improve lamb survival.

Alpacas fed calcium propionate seem to moderate their energy intake.

Alpacas have evolved digestive and metabolic adaptations that enable them to survive in environments where the available feed varies in nutritional quality. Alpacas are thought to derive glucose from the deamination of amino acids in the liver, rather than via the conversion of propionate like true ruminants. Because fibre growth is dependent on the availability of absorbed amino acids, alpacas using amino acids as a source of energy should leave less amino acids available for fibre growth. If alpacas were to obtain glucose from a source of propionate, such as calcium propionate, the dependence on amino acids would be reduced leaving more available for fibre growth. Calcium propionate was added to the ration fed to 32 alpaca wethers, and fibre production was measured to monitor important fibre attributes in response to calcium propionate. Although the diets supplemented with calcium propionate should have provided more energy than the diets without calcium propionate, the metabolisable energy intake of all animals was similar (p = 0.278). It seems that rather than sparing amino acids, the alpacas regulated their energy intake and refused to consume the additional energy offered as calcium propionate. Consequently, they produced less fibre, and the diameter of their fibre was smaller than those alpacas that were not fed calcium propionate. It seems that alpacas rely on their digestive and metabolic adaptations to efficiently obtain and conserve energy for their survival.

Follicle development, endocrine profiles and ovulation rate in adult Merino ewes: effects of early nutrition (pre- and post-natal) and supplementation with lupin grain.

In adult ewes, we tested whether ovarian function, including the response to short-term supplementation, was affected by the nutrition of their mothers during the pre-/post-natal period. A 2×2 factorial design was used with nutrition in early life (low or high) and a 6-day supplement (with or without) as factors. All ewes received three prostaglandin (PG) injections 7 days apart, and the supplement (lupin grain) was fed for 6 days from 2 days after the second until the third PG injection. We measured reproductive and metabolic hormones, studied follicle dynamics (ultrasonography), and evaluated granulosa cell numbers, aromatase activity and oestradiol (E2) concentrations in follicular fluid in healthy follicles at days 3 and 7 of supplementation. Ovulation rate was increased by 25% by exposure to high pre-/post-natal nutrition (1.5 vs 1.2; P<0.05), in association with a small decrease in FSH concentrations (P=0.06) and a small increase in insulin concentrations (P=0.07). The number of healthy antral follicles was not affected. Acute supplementation increased the number of granulosa cells (3.7±0.2 vs 3.0±0.2 million; P<0.05) in the largest follicle, and the circulating concentrations of E2 (4.6±0.3 vs 3.9±0.3 pmol/l; P<0.05) and glucose (3.4±0.03 vs 3.3±0.03 mmol/l; P<0.01). Both early life nutrition and acute supplementation appear to affect ovulation rate through changes in glucose-insulin homoeostasis that alter follicular responsiveness to FSH and therefore E2-FSH balance.

Rumen-protected methionine supplementation and fibre production in alpacas (Vicugna pacos).

Sulphur-containing amino acids are a crucial requirement for fibre production and may be supplemented in the diet of fibre-producing animals to stimulate fibre growth. The alpaca fibre industry is a developing industry in Australia with high variability in fibre production. To date, there is no evidence whether supplementing the diet of alpacas with sulphur amino acids improves fibre production. We hypothesised that supplementation with the rumen-protected sulphur amino acid, methionine would increase fibre growth in alpacas. Three groups of eight huacaya alpaca wethers were fed daily a maintenance diet supplemented with 0, 2 or 4 g of rumen-protected methionine for 7 weeks. Fibre samples were taken at the beginning and end of the study with a blood sample taken by jugular venipuncture prior to feeding on the first day of each week. Methionine supplementation had no effect on fibre diameter (p = 0.92), fibre length (p = 0.91) or fibre yield (p = 0.33). The change of season over the study affected plasma glucose (p < 0.001), plasma urea nitrogen (p < 0.001) and fibre diameter (p < 0.001). The indifference between groups may be due to the maintenance diet supplying sufficient levels of methionine, the lack of genetic potential of the experimental animals to respond to additional methionine or that the supplemental methionine was not protected in alpacas and deaminated for glucose production.

Embryo losses in sheep during short-term nutritional supplementation.

To test whether a nutritional supplement fed from 6 days before until 15 days after insemination reduces progesterone concentrations and increases embryo losses, Merino ewes were artificially inseminated (Day 0). Control ewes (n = 116) were not supplemented whereas Lupin6 ewes (n = 112) were supplemented with 500 g lupin grain daily for 6 days before insemination, and Lupin6+15 ewes (n = 122) from 6 days before until 15 days after insemination. There were no major differences between treatment groups in progesterone concentrations over the first 17 days of pregnancy. Embryo losses over Days 10-17 were lower in the Lupin6+15 than in the Control and Lupin6 groups, but the opposite occurred from Day 17-30. The concentrations of insulin and IGF-I were higher in Lupin6+15 ewes on Days 5, 12 and 17, compared with Lupin6 and Control ewes, while leptin concentrations decreased by Day 17 in the Lupin6+15 group. We conclude that feeding ewes for 15 days after mating improved embryo survival, which was associated with an increase in the concentrations of metabolic hormones and lower progesterone concentrations. However, the decrease in leptin concentrations promoted by the interruption of supplementation seems be linked to increased embryo mortality up to Day 30.

The use of a 'first-wave' model to study the effect of nutrition on ovarian follicular dynamics and ovulation rate in the sheep.

We have developed an experimental model in which groups of ewes are simultaneously experiencing the first ovarian follicular wave of their oestrous cycle. We used this 'first-wave model' in a 2 × 2 factorial experiment (ten ewes per group) to study the effect of body condition (BC) and a short-term supplement on follicular dynamics and ovulation rate. The 'first-wave' was established by giving ewes three injections of prostaglandin (PG), 7 days apart. The 6-day supplement (lupin grain) began 2 days after the second PG injection and continued until the third. Follicles were studied by ultrasound, and blood was sampled to measure glucose and hormones. The supplement increased (P<0.01) the concentrations of glucose, insulin and leptin, decreased FSH concentrations (P<0.01) and tended to increase oestradiol concentrations (P=0.06). The supplement tended to increase the number of 3 mm follicles (P=0.06). Compared with low-BC ewes, high-BC ewes had more follicular waves (P<0.05), higher concentrations of insulin, leptin and IGF1 (P<0.05) and tended to have higher FSH concentrations (P=0.09). Leptin and insulin concentrations remained high until the end of supplementation in high-BC ewes, whereas they decreased after the third day of supplementation in low-BC ewes. In conclusion, high concentrations of metabolic hormones in fat ewes are associated with the development of more follicular waves. When a supplement is superimposed on this situation, changes in glucose and metabolic hormones allow more follicles to be selected to ovulate.

A pre-partum lift in ewe nutrition from a high-energy lick or maize or by grazing Lotus uliginosus pasture, increases colostrum production and lamb survival.

This experiment tested the hypothesis that a lift in the nutrition of ewes, before lambing, to increase colostrum production would enhance lamb survival. In all, 261 mature Corriedale ewes, each with a single fetus from a synchronised mating, grazed native pasture to day 130 after mating; at which point they were weighed, condition scored and allocated to graze either native pasture or a pasture dominant with Lotus uliginosus. Five days later (14 days before the expected start of lambing) the ewes were allocated to one of four treatments and fed: (i) native pasture alone, (ii) native pasture plus a commercial high-energy lick, (iii) L. uliginosus pasture alone or (iv) L. uliginosus pasture plus whole maize. The weight, viscosity and concentration of components and immunoglobulin G in the colostrum that had accumulated at parturition, were measured for 10 ewes in each treatment. The lambs that survived to 20 days of age from the 221 ewes that were not milked, were recorded. The ewes supplemented with the lick or maize grain and those that grazed the L. uliginosus pasture alone accumulated two to three times more colostrum at birth than the ewes that grazed native pasture alone (396, 635 and 662 g v. 206 g; P < 0.01). The colostrum from the ewes that grazed only native pasture was more viscous (lower score) than that from the ewes supplemented with the lick or maize grain or the ewes that grazed the L. uliginosus pasture alone (scores of 4.1 v. 6.2, 6.5 and 6.4, P < 0.001) and, not surprisingly, the concentration of lactose in the colostrum of the ewes fed only native pasture was also much lower (1.1% v. 3.0%, 2.8% and 2.6%; P < 0.001)he survival of lambs from the ewes fed only native pasture was less than that of the lambs from ewes fed native pasture plus the commercial lick (81.8% v. 95.5%; P < 0.05) or the L. uliginosus pasture alone (92.4%, P < 0.05), and also tended to be lower than that for lambs born to ewes fed L. uliginosus pasture plus maize (91.8%, P = 0.08). The concentration of glucose in the blood of the lambs from the ewes that grazed only native pasture was lower than that of the other lambs (42.1 v. 60.2 ng/ml, P = 0.012). We conclude that the marked increase in colostrum production associated with the lift in ewe nutrition, just prior to lambing, enhanced lamb survival.

Effect of supplementation of ewes with barley or maize during the last week of pregnancy on colostrum production.

The experiment tested the hypothesis that short-term feeding of barley just before lambing would be as effective as maize in stimulating early production of colostrum. Both grains are high in starch, and should provide a substrate for lactose which, in turn, promotes lactogenesis. Thirty-five Corriedale ewes bearing single foetuses and 25 bearing twin-foetuses from a synchronised mating were fed on pasture during most of gestation. Fourteen days before the expected time of lambing the single- and twin-bearing ewes were allocated to three treatments and fed (1) a basal diet of lucerne hay to meet their nutrient requirements, (2) the basal diet plus a supplement of whole barley; or (3) the basal diet plus a supplement of cracked maize. The twin-bearing control ewes accumulated more colostrum than the single-bearing control ewes at birth (292 v. 190 g). However, supplementation with barley or maize increased the colostrum at birth to 360 and 541 g in singles and 648 and 623 g in twins. We conclude that barley is a good alternative to maize to stimulate production of colostrum especially in twin-bearing ewes whose lambs are the most likely to benefit from the supplement.

Natural methods for increasing reproductive efficiency in small ruminants.

This paper describes three strategies to improve the reproductive performance of small ruminants in ways that lead to "clean, green and ethical" animal production. The first is aimed at control of the timing of reproductive events for which we turn to the socio-sexual inputs of the "male effect" to induce synchronised ovulation in females that would otherwise be anovulatory. The second strategy, "focussed feeding", is based on our knowledge of the responses to nutrition and aims to develop short programs of nutritional supplements that are precisely timed and specifically designed for individual events in the reproductive process, such as gamete production, embryo survival, fetal programming and colostrum production. The third strategy aims to maximise offspring survival by a combination of management, nutrition and genetic selection for behavior (temperament). All of these approaches involve non-pharmacological manipulation of the endogenous control systems of the animals and complement the detailed information from ultrasound that is now becoming available. The use of such clean, green and ethical tools in the management of our animals can be cost-effective, increase productivity and, at the same time, greatly improve the image of meat and milk industries in society and the marketplace.

Nutrition and colostrum production in sheep. 1. Metabolic and hormonal responses to a high-energy supplement in the final stages of pregnancy.

We tested the hypothesis that supplementation with cracked maize during the last week of pregnancy would provide ewes with a substrate for glucose and enhance the synthesis of lactose and, consequently, their production of colostrum. Thirty single- and 30 twin-bearing ewes were fed lucerne hay and half of each group was supplemented daily with 0.75 kg per head cracked maize during the last week of pregnancy. Colostrum production and the endocrine patterns in the animals were investigated. Supplementation with maize more than doubled the mass of colostrum available at birth in unsupplemented ewes: 339 v. 145 g in single-bearing ewes and 536 v. 197 g in twin-bearing ewes (P < 0.001). The total colostrum produced in the 10 h after birth was also significantly increased by supplementation: 730 v. 475 g in single-bearing ewes and 1259 v. 631 g in twin-bearing ewes (P < 0.01). The colostrum in the supplemented ewes was also more liquid with a viscosity score of 5.8 compared with 5.7 and 4.5 in unsupplemented single- and twin-bearing ewes (P < 0.01). Supplemented ewes had higher concentrations of lactose in their colostrum at parturition (2.6% v. 1.8% in single-bearing ewes and 2.5% v. 1.4% in twin-bearing ewes; P < 0.01). The plasma concentrations of progesterone and growth hormone in supplemented ewes were lower, whereas those of IGF-I and insulin were higher, all consistent with a higher capacity to produce colostrum. It is concluded that a high-energy supplement, like maize, fed to ewes in the last week of gestation increases their capacity to produce colostrum for their lambs, particularly for ewes bearing twins.

Nutrition and colostrum production in sheep. 2. Metabolic and hormonal responses to different energy sources in the final stages of pregnancy.

Lupins and maize, with similar concentrations of metabolisable energy, should produce similar responses in colostrum production at parturition when fed during the last week of pregnancy, but, in the present study, we tested the proposal that the physical form of whole lupins would restrict intake and, therefore, the response compared with cracked lupins or maize. Fifty-five twin-bearing ewes were divided into four groups: in the last 15 days of pregnancy, 14 were fed whole lupins, 13 were fed cracked lupins, 14 were fed cracked maize and 14 received no supplement. The cracked supplements were fed in increasing amounts for 6 days to avoid acidosis. The whole lupins were fed only from Day -8. All supplementary grains increased the intake of metabolisable energy by >35%, but only ewes eating maize accumulated significantly more colostrum at parturition: control, 207 g; cracked maize, 452 g; cracked lupins, 206 g; whole lupins, 231 g (P < 0.05). Plasma urea concentrations were extremely high (approximately 10 mmol L(-1)) for both groups eating lupins and approximately double those of control ewes or those receiving maize ( P < 0.05). We conclude that gut distention is not a cause of a poor response to lupins, but the ammonia associated with near-toxic concentrations of plasma urea may be affecting the production of colostrum.