Energy Balance Nadir Research Articles

Effects of energy balance on follicular development and first ovulation in postpartum dairy cows

As milk production has increased during the past four decades, conception rates in lactating cows have declined. Although reduced reproductive performance has been associated with high milk yields, measures of postpartum ovarian activity have been more closely related to energy balance. The relationship between daily energy balance and postpartum reproductive activity is confirmed by longer intervals to first ovulation in cows with greater body condition loss. Patterns in daily energy balance, such as improvement from nadir, have been correlated with enhanced follicular function and a shorter interval to first ovulation. Such observations are consistent with increased LH pulse frequency following the energy balance nadir in lactating dairy cows. Evidence indicates a primarily hypothalamic locus for the modulation of LH secretion during negative energy balance. Formation of follicular waves after parturition begins synchronously in response to increased FSH in the first week postpartum, and is typically not a limiting factor in reproductive recrudescence. Altered follicular responsiveness to gonadotrophic support through changes in metabolic hormones such as insulin-like growth factor I (IGF-I) and insulin may contribute to impaired function of dominant follicles early postpartum. Positive relationships between changes in energy balance, peripheral IGF-I and function of dominant follicles support the identification of IGF-I and the day of the energy balance nadir as metabolic modulators of postpartum ovarian activity in dairy cows.

The Effect of Genetic Merit and Production System on Dairy Cow Fertility, Measured Using Progesterone Profiles and On-Farm Recording

The recent decline in dairy cow fertility appears to be a feature of several countries and is often linked to increased milk production, but its causes are not always obvious. A fully recorded 200-cow dairy herd, split into 2 genetic lines maintained on 2 production systems, was used to investigate the relationship between several measures of fertility, production, and energy balance. The 2 genetic lines were composed of a selection line, derived from the highest genetic merit bulls available, and a control line, maintained at the average of UK genetic merit at the time of mating. The production systems were a high-concentrate and a high-forage system. Thrice-weekly milk progesterone samples allowed an objective measurement of luteal cycling activity, and farm observations of estrus, services, and calving provided data on various measures of fertility. Energy balance in early lactation was calculated from daily live weight and weekly BCS measurements. Control line cows commenced luteal activity (C-LA) 6 d before selection line cows, had their first heat 14 d earlier, and had longer gestation periods by 3.7 d. They also had a lower incidence of silent heats. Cows on the high-forage system commenced luteal activity 6 d before those on the high-concentrate system, had longer gestation intervals by 3.9 d, held to first service better, had longer luteal phases and shorter interluteal periods in their estrus cycles. Characteristics of energy balance were used to see if they could account for the fertility differences between both genetic lines and systems. The commencement of luteal activity and day of first heat were analyzed using a REML mixed model approach. Mean energy content and mean energy balance over the first 25 d of lactation had an effect on C-LA and accounted for the differences found between production systems but not genetic lines. Day of energy balance nadir, mean energy content in the first 25 d, and C-LA affected day of first heat, but the differences between genetic lines were still apparent. These results suggest a link between high performance and reduced dairy cow fertility; high performance originating from different feeding systems was largely due to differences in energy balance, whereas those originating from genetics remained when energy balance characteristics were taken into account. This suggests a real genetic change in fertility due to selection for high genetic merit.

A comparison of energy balance and metabolic profiles of the New Zealand and North American strains of Holstein Friesian dairy cow

The milk production, energy balance (EB), endocrine and metabolite profiles of 10 New Zealand Holstein Friesian (NZ) cows and 10 North American Holstein Friesian (NA) cows were compared. The NA cows had greater peak milk yields and total lactation milk yields (7387 v. 6208 kg; s.e.d. = 359), lower milk fat and similar protein concentrations compared with the NZ cows. Body weight (BW) was greater for NA cows compared with NZ cows throughout lactation (596 v. 544 kg; s.e.d. = 15.5), while body condition score (BCS) tended to be lower. The NA strain tended to have greater dry matter intake (DMI) (17.2 v. 15.7 kg/day; s.e.d. = 0.78) for week 1 to 20 of lactation, though DMI as a proportion of metabolic BW was similar for both strains. No differences were observed between the strains in the timing and magnitude of the EB nadir, interval to neutral EB, or mean daily EB for week 1 to 20 of lactation. Plasma concentrations of glucose and insulin were greater for NA cows during the transition period (day 14 prepartum to day 28 postpartum). Plasma IGF-I concentrations were similar for the strains at this time, but NZ cows had greater plasma IGF-I concentration from day 29 to day 100 of lactation, despite similar calculated EB. In conclusion, the results of this study do not support the premise that the NZ strain has a more favourable metabolic status during the transition period. The results, however, indicate that NZ cows begin to partition nutrients towards body reserves during mid-lactation, whereas NA cows continue to partition nutrients to milk production.

Strategies to improve fertility of high yielding dairy farms: Management of the dry period

Reproductive performance of dairy cattle has been related to a wide variety of indicators of energy status, e.g., extent of negative energy balance, time of energy balance nadir, body weight loss, body condition score, and body condition score loss. Energy balance begins to decrease during the last few weeks prior to calving primarily due to a 30–35% reduction in feed intake. Cows typically remain in negative energy balance for five to seven weeks postpartum. Nutritional strategies to improve energy balance during the transition period include fat supplementation and feeding additional nonfiber carbohydrate. Unfortunately, neither approach is likely to markedly enhance energy status, although fat supplementation may increase reproductive efficiency independent of any effect on energy balance. Alternative management strategies may be required to improve fertility of dairy cows. Shortening or eliminating the dry period may improve energy status of dairy cows and increase reproductive efficiency. Shortening or eliminating the dry period may enhance dry matter intake during the transition period, decrease milk energy output, or both. A preliminary study using small animal numbers indicated that reducing dry period length to 28 or 0 days may decreases days to first ovulation, increase first service conception rate, and decrease days open. A follow-up study employing large animal numbers confirmed that reducing dry period length from 55 to 34 days can decrease days to first ovulation and decrease the percentage of anovular cows. The reduction in days open was greater for older cows than second parity cows. The reduction in days open was not related to effects of treatment on milk yield. Shortening or eliminating the dry period may be a more successful approach to improving reproductive efficiency than diet manipulation.

The postpartum uterus.

The cow is notable among the domestic species for the almost ubiquitous bacterial contamination of the uterine lumen after parturition and the high incidence of clinical uterine disease. Such uterine disease not only disrupts uterine tissues, but also ovarian follicle growth and function by perturbation of the hypothalamus, pituitary, and ovary. Other events such as the postpartum negative energy balance also have similar multilevel effects on postpartum endocrinology. Return of ovarian cyclical activity is dependent on the innate immune system resolving uterine bacterial contamination,prompt uterine involution, and a short interval to the negative energy balance nadir. Although the risk factors for uterine disease have been described, preventive strategies are not widely adopted. Thus, veterinarians must identify and treat uterine disease efficiently to limit their negative effect on fertility.

Postpartum Reproductive Function: Association with Energy, Metabolic and Endocrine Status in High Yielding Dairy Cows

The effect of energetic, metabolic and endocrine status upon onset of ovarian cyclicity as well as time empty and conception at first service was assessed in 90 multiparous Holstein cows (mean 9'500 kg energy-corrected milk/305 d) in a research farm in Switzerland. Dry matter intake, milk yield and body weight were measured daily from week 2 antepartum (a.p.) through wk 20 postpartum (p.p.). Milk composition was determined 4 times/wk and milk acetone was measured weekly. Blood samples for the determination of glucose, non-esterified fatty acids (NEFA), cholesterol, creatinine, albumin, urea, s-hydroxybutyrate (BHBA), leptin, insulin, insulin-like growth factor-1 (IGF-1), growth hormone, 3,5,3'-triiodothyronine (T 3 ) and thyroxine (T 4 ) concentrations, and of activities of aspartate amino transferase (ASAT), lactate dehydrogenase (LDH) and glutamate dehydrogenase (GLDH) were taken 2 wk a.p. and in wk 1-16 and 20 p.p. between 0730-0900. Cows were gynaecologically examined weekly. The onset of first ovulation was specified by skim milk progesterone determination by a radioimmunoassay (twice/wk) and by rectal palpation. Energy balance (EB) traits were calculated and expressed as accumulated negative EB from parturition to EB equilibrium (AUC), EB nadir (EBN), rate of EB recovery following EBN (EBR), time from parturition to EBN (DEBN), and time from parturition to EB equilibrium (DEBE). Results: Resumption of ovarian cyclicity postpartum was not reflected by any of the EB traits. However, a small degree of EBN and a fast EBR were related to a short period from parturition to conception. Survival analysis with timevarying covariates revealed that high plasma concentrations of T 3 and T 4 p.p. were significantly associated with a high chance of an early start of the ovarian cycle, and high concentrations of glucose and cholesterol were associated with a short time empty, respectively. Conception at first service was positively associated with EB at first service and with progesterone concentration at 10-13 d after first service. However, leptin, as most other metabolic and endocrine traits, was not associated with any of the reproductive traits. In conclusion, thyroid hormones may play an important role in the onset of the ovarian cycle p.p., and a good energy status enhances the chance of conception at first service and shortens time empty.

Postpartum reproductive function: association with energy, metabolic and endocrine status in high yielding dairy cows

This study evaluated the effect of metabolic, endocrine and energy status on onset of ovarian cycle, days open (DO), and conception at first service in 90 multiparous Holstein cows, housed at a research farm. Dry matter intake, milk yield and body weight were measured daily from Week 2 antepartum (a.p.) to Week 20 postpartum (p.p.). Milk composition was determined four times per week and milk acetone was measured weekly. Blood samples for the determination of glucose, non-esterified fatty acids, cholesterol, creatinine, albumin, urea, β-hydroxybutyrate, leptin, insulin, insulin-like growth factor-1, growth hormone, 3,5,3′-triiodothyronine (T3), and thyroxine (T4) were taken 2 weeks a.p., in Weeks 1–16, and 20 p.p. between 7:30 and 9:00h. The onset of ovarian cycle was specified by weekly gynecological examination and by skim milk progesterone determination by radioimmunoassay (twice per week). Energy balance (EB) traits were calculated and expressed as accumulated negative EB from calving to EB equilibrium, EB nadir (EBN), rate of EB recovery after EBN (EBR), and time from calving to EBN and to EB equilibrium, respectively. The onset of ovarian cycle p.p. was not related to EB. However, a low degree of EBN and a fast EBR were associated with fewer DO, and EB at first service was positively related to conception. High plasma levels of T3 and T4 p.p. were associated with an early start of ovarian cycle, and high concentrations of glucose and cholesterol with a short calving to conception interval. Conception at first service was positively related to EB at first service and progesterone concentration 10–13 days after first service. In conclusion, thyroid hormones may play an important role in resumption of ovarian cyclicity p.p., and a good energy status enhances the chance of conception at first service and shortens DO.

Energy Balance of Dairy Cattle in Relation to Milk Production Variables and Fertility

Variables derived from milk yield records were investigated to find an easy to measure and readily available indicator of the energy balance status of a lactating cow. Weekly energy balances during the first 180 d in milk (DIM) were calculated from weekly yield, live weight, and energy intake records for 470 first lactation heifers. The energy balance curve for each cow was estimated using a random regression model. From each curve, three measures were calculated to describe the energy balance status: 1) total energy deficit in early lactation, 2) interval for return to positive energy balance, and 3) lowest value (nadir) for energy balance. Mean energy deficit per lactation was 776.8 MJ of NEL/d, interval for return to positive energy balance was 41.47 d, and nadir was –33.72 MJ of NEL/d. Regression analysis to relate these variables to interval to start of luteal activity (measured using progesterone profiles) showed that a low nadir of energy balance was related to delayed resumption of luteal activity. In general, a 10 MJ of NEL/d lower nadir of energy balance corresponded to a delay of ovulation of 1.25 d. A relatively strong decrease in fat percentage during early lactation was significantly correlated with lower nadir of energy balance, larger energy deficit, and later return to positive energy balance. The maximal correlation was between nadir of energy balance and a decrease of milk fat percentage. This correlation remained above 0.60 throughout the first 26 DIM but dropped to 0.14 at 180 DIM. Large decreases in milk fat percentage were related to high initial fat percentages at the start of lactation and slightly lower fat percentages later during lactation. Hence, we concluded that a decrease in fat percentage during early lactation might serve as an indicator of energy balance.

Modeling of energy balance in early lactation and the effect of energy deficits in early lactation on first detected estrus postpartum in dairy cows.

The general pattern of energy balance in early lactation was modeled. For this purpose, several lactation curves were investigated. The best fitting curve was fit in a random regression model that provided predicted energy balance curves for all lactations. By use of these curves, total energy deficit per lactation, postpartum interval of return to positive energy balance, and nadir of energy balance was determined. In predicted energy balances, nadir of energy balance was present, but variation in postpartum interval to nadir was small. First lactation cows had a smaller energy deficit in early lactation than did older cows. Differences among cows in nadir and total energy deficits in early lactation were large. Both were related to the postpartum interval to first detected estrus with a larger energy deficit and a smaller nadir corresponding to a larger postpartum interval to first detected estrus. From this study, it can be concluded that large energy deficits in early lactation delay first detected estrus.

Energy Balance, Metabolic Hormones, and Early Postpartum Follicular Development in Dairy Cows Fed Prilled Lipid

The objectives of this study were to relate energy balance and metabolic hormones during the early postpartum period in dairy cows with dominant follicle development before first ovulation and to evaluate the effects of prilled lipid on follicular development during the first follicular wave after parturition and the postpartum anovulatory interval. At parturition, 42 cows received a control diet (4.8% fat) or a diet supplemented with prilled fatty acids (7.0% fat). Energy balance was determined daily. Ovarian follicular development was monitored by ultrasonography, and blood plasma or serum was analyzed for estradiol, progesterone, and metabolic hormones. Dry matter intake was lower in cows supplemented with dietary lipids during the first 4wk of lactation, but energy intake, energy balance, and the postpartum anovulatory interval were similar between diets. A wave of follicular development occurred in all cows during the 2nd wk postpartum, and 50% of all cows ovulated their first dominant follicle. Numbers of follicles that were 3 to 5mm, 6 to 9mm, and 10 to 15mm on d 8 postpartum were similar between diets and unrelated to energy balance or metabolic hormones. Diameter of the dominant follicle during d 8 to 14 postpartum and maximum diameter of the first-wave ovulatory follicle did not differ between diets. Cows with nonovulatory first-wave dominant follicles had lower mean plasma concentrations of estradiol during d 8 to 14 postpartum, a longer interval to the day of the energy balance nadir, lower serum concentrations of IGF-I, and higher 4% FCM yield than did cows with ovulatory first-wave dominant follicles. Serum IGF-I during d 1 to 13 was positively correlated with plasma estradiol during d 8 to 14 postpartum. Possibly because of reductions in dry matter intake, the consumption of prilled lipid by dairy cows during early lactation may be ineffective in altering energy balance, follicular development, and the postpartum anovulatory interval. Ovulation failure of dominant follicles early in the postpartum period is associated with greater production of 4% fat-corrected milk, a delayed energy balance nadir, and reduced concentrations of peripheral IGF-I.

Energy balance and ovarian follicle development prior to the first ovulation postpartum in dairy cows receiving three levels of dietary fat.

The objectives of this study were to characterize early postpartum (PP) follicular development in dairy cows and to evaluate the influence of energy balance (EB) and the level of dietary fat on dominant follicle development and function. Forty-five multiparous Holstein cows were fed either low (LF, 3.3%), moderate (MF, 5.2%), or high (HF, 7.1%) total dietary fat beginning at parturition, and daily EB was determined for all cows. Follicular development was monitored by ultrasonography, and blood plasma was analyzed for metabolites, metabolic hormones, progesterone, estradiol, and FSH. After an increase (p < 0.01) in mean plasma FSH during Days 1-5, all cows experienced a wave of follicular development during the second week PP regardless of diet or EB. Feeding the MF (p < 0.05) or HF (p < 0.06) diet resulted in a greater number of class IV (> 15 mm) follicles on Day 14 PP as compared to the LF diet. Cows fed the MF diet had higher (p < 0.05) peak plasma estradiol during the first follicular wave and a shorter (p < 0.05) interval to first ovulation than both LF and HF cows. Follicular development prior to first ovulation was characterized by either ovulation of the first dominant follicle (OV, n = 19), one or more waves of nonovulatory dominant follicles (NOV, n = 18), or the formation of a follicular cyst (CYST, n = 8). Ovulation failure during the first follicular wave PP was accompanied by lower (p < 0.001) peak plasma estradiol (0.96 +/- 0.36 vs. 5.0 +/- 0.35 pg/ml), a smaller (p < 0.01) maximum follicle diameter (16.0 +/- 0.9 vs. 20.0 +/- 1.0 mm), lower (p < 0.09) levels of plasma insulin-like growth factor-I (IGF-I; 73.5 +/- 10.1 vs. 102.9 +/- 9.9 ng/ml), and a longer (p < 0.02) interval to the EB nadir (14.6 +/- 1.9 vs. 6.4 +/- 1.9 days). The number of days to first ovulation was positively correlated with days to the EB nadir (r = 0.55), with dominant follicles that emerged after the EB nadir exhibiting enhanced (p < 0.01) production of estradiol and greater (p < 0.01) ovulatory success. The development of large (> 10 mm) dominant follicles was not a limiting factor in PP reproductive recovery, and moderate dietary fat shortened the interval to first ovulation. Follicular competence early PP was associated with higher plasma IGF-I and a shorter interval to the EB nadir.

Energy balance, first ovulation and the effects of naloxone on LH secretion in early postpartum dairy cows.

Relationship among energy balance (EB), first ovulation, circulating metabolites and opioid involvement in peripheral LH concentrations were assessed in 40 multiparous Holstein cows assigned randomly to an experiment with a 2 x 2 factorial arrangement of treatments. Cows were either milked normally (2x/d) and expected to be in negative EB (LAC) or were milked for only 1 d postpartum (PP) to ensure a rapid return to positive EB (NONLAC). Blood samples were collected from NONLAC cows every 12 min for 16 h on d 7, 10, and 14 PP and from LAC cows on d 7, 14, 21 and 28 PP. Within each group, half the cows received naloxone (50 mg/h) and half received saline during h 9 to 16 of each frequent sampling series. The mean number of LH pulses/8 h for NONLAC cows was 4.3, 4.7 and 5.3 at 7, 10 and 14 d PP and for LAC cows 4.3, 5.7, 6.4 and 7.0 at 7, 14, 21 and 28 d PP, respectively. The LH pulse frequency was not different between NONLAC and LAC cows at 7 and 14 d PP, yet NONLAC cows had fewer days to first ovulation (P less than .01). Naloxone did not affect any parameters of LH secretion in either group. Following parturition, NONLAC cows averaged 4.0 d to negative EB nadir and 14.3 d to first ovulation. The LAC cows averaged 13.6 d to negative EB nadir and 27.0 d to first ovulation. Days PP to first ovulation were highly correlated (r = .85) with days PP to negative EB nadir.(ABSTRACT TRUNCATED AT 250 WORDS)

Effects of Excess Degradable Protein on Postpartum Reproduction and Energy Balance in Dairy Cattle

The objectives of this study were to observe endocrine and reproductive responses of cows and heifers fed two diets (16 and 19% CP), which met undegradable protein requirements but differed in rumen degradable protein. Cows (n = 33) and heifers (n = 32) were randomly assigned within parity to diets at calving and remained on diets for 20 d after first breeding. Energy balance was determined twice weekly through the first luteal phase. Blood and milk samples were taken three times per week. Diet did not affect average daily energy balance, days to negative energy balance nadir, days to first ovulation, days to first service, or plasma glucose concentrations. First service conception rate was lower (31% vs. 48%) and plasma urea higher in animals fed the high protein diet. Days to energy balance nadir was correlated with days (r = .75) to first ovulation. Luteinizing hormone pulse frequency increased and pulse amplitude decreased in frequent samples (12-min intervals for 8h) collected at 14 d postpartum versus sampling after the energy balance nadir. These data suggest that energy balance status plays an important role in determining the postpartum return of cyclic ovarian activity. Feeding excess CP as rumen degradable protein elevated plasma urea concentrations and decreased first service conception rate.

Energy balance and pulsatile LH secretion in early postpartum dairy cattle

The relationships between energy balance (EB), pulsatile LH secretion and circulating levels of insulin and various energy metabolites were assessed in ten lactating Holstein cows. Cows were blood sampled every 12 min via indwelling jugular catheters for 8 hr twice weekly beginning in the first week postpartum (PP) through first ovulation (1 stOV). Days to negative EB nadir and days to 1 stOV were highly correlated (P < .02). LH pulse frequency (P < .01), LH baseline (P < .05) and mean LH (P < .1) increased, while LH pulse amplitude tended to decrease when comparing the frequent sampling series immediately before and after the negative EB nadir. Plasma levels of non-esterified fatty acids (NEFA, r = −.40, P < .01), insulin (r = .38, P < .01), glucose (r = .31, P < .01) and beta-hydroxybutyrate (r = .19, P < .06) were correlated with EB. For within animal comparisons, NEFA was significantly correlated (r = −.44 to −.89, P < .05) with EB in 8 of 10 cows, but no other correlations were significant. Additionally, none of the metabolites nor insulin were significant in multiple regression models for 1 stOV. Therefore, it appears that pulsatile LH secretion is suppressed until the negative EB nadir is reached, at which time LH pulse frequency increases stimulating 1 stOV. Since NEFA and EB are directly related, NEFA may serve as a peripheral signal of EB to the central nervous system.

Association Between Energy Balance and Luteal Function in Lactating Dairy Cows

Our objective was to determine the relationship between energy balance and secretion of progesterone in lactating dairy cows. Eight primiparous and 24 multiparous lactating Holstein cows were studied from parturition to 100 d postpartum or conception. Cows calved normally and remained healthy throughout the study. All cows were fed ad libitum a total mixed diet formulated to satisfy requirements for maintenance and lactation. Intake of feed and production of milk per cow were measured twice daily. Body weight was determined weekly. Daily energy balance was determined by subtracting energy required for maintenance and lactation from intake of energy. Concentrations of progesterone were determined in milk sampled every 3rd d. For at least 4 successive d postpartum, 81% of cows were in negative energy balance. Variation in energy balance was explained largely by intake of energy. Duration of luteal phases was not associated with energy balance. Energy balance within 9 d postpartum was correlated positively with concentration of progesterone within second and third postpartum luteal phase. Postpartum interval to nadir and magnitude of nadir of energy balance interacted to reduce progesterone within second and third postpartum estrous cycles. Thus, in lactating cows, secretion of progesterone is reduced by spontaneous caloric deficit and is modulated by timing and magnitude of maximal caloric deficit. Spontaneous caloric deficit is a potential source of infertility in lactating dairy cows.