Estimation of genetic parameters among reproductive and growth traits in yearling heifers

Growth and reproductive data were obtained on 779 beef heifers at the San Juan Basin Research Center, Hesperus, Co. Genetic parameters were estimated for age of puberty (AOP), age of first calving (AOC), julian day of first calving (DOC), julian day of second calving (DOSC), birth weight, weaning weight, yearling weight, and average daily gain from weaning to yearling and to cycling weights. The least squares model included birth year, age of dam and breed as fixed effects, sire/breed as a random variable, and day of birth and percent inbreeding as covariates. Day of birth was not included in the analyses of AOC, DOC or DOSC. Paternal half-sib estimates of heritability were: AOP, .10 +/- .17; AOC, .01 +/- .12; DOC, .09 +/- .13 and DOSC, .36 +/- .18. Genetic and phenotypic correlations were generally favorable, but genetic correlations were variable with large standard errors. Inbreeding had a detrimental effect on reproductive traits, and a seasonal effect was present for AOP.     

Calving date versus calving interval as a reproductive measure in beef cattle

Calving date and calving interval records obtained from the Beef Cattle Division of Pioneer Hi-Bred International, Des Moines, Iowa, were studied to determine the reproductive measure of choice when a fixed breeding season is employed. Fixed model, least-squares procedures were used to analyze 2,411 records collected on purebred Red Angus, Angus and Hereford cows during the years 1969 through 1976. Calving interval decreased .86 d and calving date was delayed .11 d for each 1-d delay in previous calving date. A 1-d increase in gestation length was associated with a 1.17-d delay in calving date and 1.17-d increase in calving interval. The bias caused by the use of a fixed breeding season was determined to be much greater for calving interval than for calving date, as evidenced by the strong dependence of calving interval on previous calving date. Because calving date is likely to be more heritable and has clear economic significance, and because selection for earlier calving date should not lead to adverse response in other reproductive traits, calving date was preferred over calving interval as a reproductive measure.     

三河牛繁殖性状遗传参数的评估

本研究以1986年中国自主培育兼用牛新品种--三河牛,在内蒙古海拉尔谢尔塔拉种牛场核心群5 257头1998-2012年20 949条繁殖记录为研究材料,以青年牛首次妊娠日龄、青年牛首次产犊日龄、成母牛妊娠期、成母牛空怀期、产犊间隔为研究对象,用SAS 9.13、DMU软件对数据进行处理,采用AI-REML结合EM算法并配合多性状动物模型对各性状影响因素方差组分进行估计,估算出各性状遗传力,并利用各性状育种值分析其遗传趋势.结果显示,青年牛首次妊娠日龄、青年牛首次产犊日龄、成母牛妊娠期、成母牛空怀期、产犊间隔遗传力分别为0.0552、0.0638、0.0527、0.1096、0.0844,繁殖性状除成母牛空怀期遗传力为0.1096外,其余均小于0.1,属于低遗传力性状.青年牛首次妊娠日龄、青年牛首次产犊日龄、成母牛妊娠期、成母牛空怀期、产犊间隔育种值遗传趋势总体上无明显下降趋势,三河牛繁殖性能保持良好.该试验结果为三河牛优化育种方案、提高选种准确性提供重要理论依据.     

Simulation of genetic control of reproduction in beef cows. II. Derived genetic parameters

A stochastic computer simulation model was used to predict heritability (h2) and repeatability (t) of derived reproductive traits in beef cattle as a function of underlying, normally-distributed genetic variation in interval from calving to first estrus (postpartum interval) and single-service conception rate. Traits simulated were mating rate, first-service conception rate, overall conception rate, date of first service, number of services during a 63-d breeding season, calving date and postpartum interval. Weight of calf weaned was also simulated on a per-cow-exposed, per-cow-mating, per-cow-calving, per-calf-weaned or adjusted 205-d basis. Derived estimates of h2 for mating rate and date of first service were not significantly different from 0, reflecting the high proportion of cows that were predicted to cycle within the first 21 d of breeding. Estimates of h2 were significant and approached the level of input h2 for overall and first-service conception rate and number of services, suggesting that these traits may be potential selection criteria. Values of h2 for calving date were significant but considerably smaller than input h2. Estimates of t were much larger than h2 for date of first service and calving date, even when no nonadditive genetic or permanent environmental effects were explicitly simulated. Estimates of h2 for weaning weight per cow exposed and especially for weaning weight per cow calving (in the presence of random calf death losses) were much lower than underlying h2 values. Culling of open cows generally reduced genetic variances of derived reproductive traits to negligible levels within three calf crops.     

Genetic Evaluation of Reproductive Traits in Sanhe Cattle

This study was conducted on 20 949 reproductive records from 1998 to 2012 of 5 257 Sanhe cattle in Xiertala cattle farm,Inner Mongolia,which was a synthetic breed formed in China in 1986.Age at first pregnancy in heifer (AFPH),age at first calving in heifer (AFCH),gestation length in cow (GLC),days open in cow (DOC),and calving interval in cow (CIC) were considered for genetic evaluation.SAS 9.13 and DMU software were used for data processing,and AI-REML combined EM algorithm based on multiple traits animal model was employed for estimating variance components.The heritability for each trait were then calculated,and breeding value was used to analyze the genetic trends.The results showed that the estimated heritabilities of age at first pregnancy in heifer,age at first calving in heifer,gestation length in cow,days open in cow,calving interval in cow were 0.0552,0.0638,0.0527,0.1096 and 0.0844,respectively.The heritabilities were all less than 0.1 except days open (0.1096),indicating these were low inheritable traits.In general,trends of EBVs for each trait didn't show any defined progresses and indicating good reproductive performance maintained in Sanhe cattle.These results lay a theoretical foundation for optimizing breeding programs and improving the accuracy of selection in Sanhe cattle.     

Genetic parameters and genetic trends for age at first calving and calving interval in an Angus-Blanco Orejinegro-Zebu multibreed cattle population in Colombia

Genetic parameters and genetic trends for age at first calving (AFC), interval between first and second calving (CI1), and interval between second and third calving (CI2) were estimated in a Colombian beef cattle population composed of Angus, Blanco Orejinegro, and Zebu straightbred and crossbred animals. Data were analyzed using a multiple trait mixed model procedures. Estimates of variance components and genetic parameters were obtained by Restricted Maximum Likelihood. The 3-trait model included the fixed effects of contemporary group (year-season of calving-sex of calf; sex of calf for CI1 and CI2 only), age at calving (CI1 and CI2 only), breed genetic effects (as a function of breed fractions of cows), and individual heterosis (as a function of cow heterozygosity). Random effects for AFC, CI1, and CI2 were cow and residual. Program AIREMLF90 was used to perform computations. Estimates of heritabilities for additive genetic effects were 0.15 ± 0.13 for AFC, 0.11 ± 0.06 for CI1, and 0.18 ± 0.11 for CI2. Low heritabilities suggested that nutrition and reproductive management should be improved to allow fuller expressions of these traits. The correlations between additive genetic effects for AFC and CI1 (0.33 ± 0.41) and for AFC and CI2 (0.40 ± 0.36) were moderate and favorable, suggesting that selection of heifers for AFC would also improve calving interval. Trends were negative for predicted cow yearly means for AFC, CI1, and CI2 from 1989 to 2004. The steepest negative trend was for cow AFC means likely due to the introduction of Angus and Blanco Orejinegro cattle into this population.     

Effects of breeding season length and calving season on range beef cow productivity

A 5-yr study was conducted beginning in 1983 with 460 cows to evaluate the effects of three breeding seasons (30, 45, and 70 d in length) and two times of spring calving, March (early) and April (late), on cattle production under Nebraska Sandhills range conditions. Criteria evaluated included pregnancy and weaning percentages, calving date and distribution, cow weights and body condition at four intervals, calf birth and weaning weights, and cow productivity. The 30-d breeding season included a 10-d estrus synchronization and AI period; in the other breeding seasons only natural breeding was used. The same sires were used over the entire study period. Percentage of cows pregnant and percentage of calves weaned were lower (P less than .01) for cows bred for 30 d than for cows bred for 45 or 70 d. Average calving dates were similar among the breeding groups within the early and late calving herds. Pregnancy rates from AI were higher (P less than .01) for the cows calving in April (64%) than for the cows calving in March (41%). Cows calving in April lost less weight between precalving and prebreeding and were heavier (P less than .05) at prebreeding time than the cows calving in March. Calf weaning weights were not different (P greater than .10) among any of the breeding season groups or between the two calving herds when calves were weaned at a similar age. Cow productivity (calf weaning weight per breeding female) was highest (P less than .05) for the cows bred for 70 d (186 kg), intermediate for the cows bred for 45 d (172 kg), and lowest for cows bred for 30 d (162 kg).(ABSTRACT TRUNCATED AT 250 WORDS)     

The Estimation of Genetic Parameters of Holstein Dairy Cows in Ningxia Region

The genetic parameters of milk fat percentage (FP), milk protein percentage (PP), somatic cell score (SCS), milk yield of 305 days (MYD), age at first calving (AFC),calving interval (CI) and linear classification scores (LCS) were estimated using the DHI data of 2008 to 2016 in Ningxia area of 27 444 Holstein dairy cows and type records. With the aid of DMU v 6.0 AI-REML software, DMU combined with EM algorithm and multi character animal model was used to model the influence factors of the birth season, herd, year, parity and individual additive genetic effect. The results showed that FP, PP, SCS, MYD, AFC, CI and LCS's heritability were 0.14, 0.19, 0.19, 0.31, 0.37, 0.10, 0.07, respectively. At the same time, the breeding value, genetic correlation and CPI2 values of different traits were calculated. The genetic evaluation for dairy cows in Ningxia, could be more in-depth understanding of the basic situation of the herd, to provide basic data for the construction of dairy cow breeding matching selection, planning and selection index.     

宁夏地区荷斯坦奶牛遗传参数估计

利用2008-2016年出生的27 444头宁夏地区荷斯坦奶牛生产性能（dairy herd improvement,DHI）数据及体型线性评分数据计算乳脂率（fat percentage,FP）、乳蛋白率（protein percentage,PP）、体细胞评分（somatic cell score,SCS）、305 d产奶量（milk yield of 305 days,MYD）、初产日龄（age at first calving,AFC）、产犊间隔（calving interval,CI）和体况评分（linear classification scores,LCS）的遗传参数。利用DMU v 6.0软件,采用AI-REML模块结合EM算法并配合多性状动物模型,以季节、场、年份、胎次和动物个体加性遗传效应作为模型的影响因素。计算结果表明,宁夏地区奶牛的FP、PP、SCS、MYD、AFC、CI和LCS的遗传力分别为0.14、0.19、0.19、0.31、0.37、0.10和0.07,同时得出了奶牛不同性状的育种值、遗传相关和奶牛生产性能指数2（China performance index 2,CPI2）值。本研究通过对宁夏地区奶牛DHI数据的深挖和遗传评估,准确地把握了宁夏奶牛群体结构特征,对于奶牛的选种选配、育种规划和选择指数的构建有重要意义。     

Maternal heterosis and grandmaternal effects in beef cattle: preweaning traits

Crossbred progeny from females, 3 yr old and older, representing Brown Swiss (B), Red Poll (R), Hereford (H) and Angus (A) maternal grandsires and maternal granddams in a diallel crossing experiment were evaluated to estimate breed mean maternal heterosis (mean hMi) for each breed, maternal heterosis for specific breed cross females (hMij), average maternal heterosis for all crosses (mean hM) and breed grandmaternal effects (gM'i) for preweaning calf traits. Estimates of mean hM were significant for birth date and weight, 200-d weight and 200-d weight/cow exposed. Even though calves with crossbred dams weighed more at birth, they did not differ from calves with straightbred dams in frequency of calving assistance. The estimates of hMij were similar for most crosses and most traits, except 200-d weight and 200-d weight/cow exposed. The largest heterotic advantage was exhibited by progeny of BH reciprocal cross females, which exceeded crossbred progeny from BB and HH females for 200-d weight/cow exposed by 35.9 kg. The smallest heterotic effect was exhibited by crossbred progeny of RA reciprocal cross females. Progeny with R maternal granddams exhibited a higher (P less than .05) live calf born and weaned percentage than progeny with H maternal granddams. Progeny with B maternal granddams were born later (P less than .05) in the calving season than progeny with R maternal granddams. Progeny with A maternal granddams exhibited a lower frequency of calving assistance than progeny with H maternal granddams, and progeny with H maternal granddams were heavier (P less than .05) at 200 d than progeny with R maternal granddams. There was a tendency for estimates of breed grandmaternal effects (gMi) to exhibit an inverse relationship with estimates of breed maternal effects (gMi).     

中国荷斯坦牛青年初产年龄和成年母牛产犊间隔遗传参数估计

本研究旨在估计河南省青年牛初次产犊年龄与成年母牛产犊间隔遗传参数。收集河南省17个牧场2008—2017年14406头青年牛的初次产犊年龄记录与8912头成年母牛的产犊间隔记录,使用DMU软件DMU_AI模块采用AI-REML结合EM算法的动物模型估计性状遗传参数。动物模型中的固定效应包括出生年季、产犊年季和牧场效应,产犊间隔性状动物模型还包括胎次效应;随机效应包括加性效应和残差效应。结果表明:青年牛初次产犊年龄的遗传力为0.21、成年母牛产犊间隔性状遗传力为0.05,与相似研究结果相近,为制定区域性、精细化的育种目标提供理论参考。     

Parameters to monitor dairy herd fertility and their relation to financial loss from reproductive failure

Relationships between herd fertility measurements and financial loss from reproductive failure in dairy herds were studied. Financial losses attributable to prolonged calving interval and forced replacement from reproductive failure were considered. Herd fertility parameters were calculated from artificial insemination and calving data (i.e. calving to first service interval, non-return rate 56 days after first service, percentage of correct inseminations carried out in the interval 18–24 days, fertility status, calving interval, an oestrus index and number of insemination per average cow present in the herd.

The herd fertility parameters were moderately-highly related to loss due to suboptimal calving interval (r=0.20−0.79 in absolute values), but only slightly related to losses due to forced replacement (r<0.17 in absolute values).

Repeatabilities, calculated over a 3-year period, were high for the interval to first service, non-return rate and the oestrus index (0.52−0.67) and moderate for percentage correct reinseminations, fertility status, calving interval and loss due to suboptimal calving interval (0.38−0.48). Repeatability of loss due to forced replacement was low (0.20).

In a regression analysis no herd fertility parameter was fitted with respect to loss from forced replacement. Loss due to suboptimal calving interval at herd level was best estimated by the oestrus index (R²=0.63), the addition of the interval to first service to the regression equation explained a further 10% of the variation between herds. It is suggested that the oestrus index and the interval to first service should be presented as management aids to monitor herd fertility.     

Post-partum Reproductive Performance of Sanga Cattle in Smallholder Peri-urban Dairy Herds in the Accra Plains of Ghana

The reproductive performance of 76 Sanga cows was studied from February 1995 to July 1996 in smallholder peri-urban dairy systems in the coastal savanna zone of the Accra plains of Ghana. The interval from calving to resumption of cyclic ovarian activity was 101±7 days; that from calving to conception was 152±4 days; gestation length was 292±16 days; and the calving interval was 444±16 days. The effects of location, season of calving, parity, body condition score and sex of calf were studied. Calving to cycling interval was significantly shorter in the dry season than in the wet season, and decreased significantly with increasing body condition score. Calving to conception interval was affected only by location. Gestation length was not affected by any of the variables studied. Calving interval was affected by location, season of calving, parity and body condition score. It is concluded that long post-partum anoestrus leads to prolonged calving intervals and poor reproductive performance. Feed supplementation to improve the nutritional status and body condition score of cows and early weaning of calves could be introduced to enhance reproductive performance.     

Some aspects of reproductive performance in selected town-supply dairy herds

Breeding and production data were collected from 20 town-supply dairy herds in the Manawatu/Horowhenua region. Calving interval, calving to first service interval, first service to conception interval, inter-service interval, calving rate to first service and services per cow calving were 384, 85, 11, 43 days, 51% and 1.7, respectively, for the pooled population of 12, 056 calvings. The mean milk yield was 3730 litres per cow and the mean lactation length 291 days. Age, herd, season within year, and year, had statistically significant effects on reproductive performance, although the amount of variation due to these variables was small. The younger cows (2-year-olds) were relatively poor performers. Reproductive performance was best during the spring. Days from calving to first service, and from first service to conception, were of similar importance in determining differences in calving interval. A high incidence (25%)of return intervals of more than 49 days was found between first and second service. It was concluded from the data that a programme for breeding each cow as she comes into oestrus 46–66 days post-partum is a simple and appropriate procedure for achieving a 365-day calving interval. Such a programme is more likely to be successful if continued attention is paid to both heat-detection procedures and the nutritional requirements of animals during rearing, late pregnancy and early lactation, and if routine pregnancy testing 6 to 8 weeks after service is used to recognise the problems of post-service anoestrus.     

Genetic parameters for a maternal breeding goal in beef production

New maternal breeding values have been developed for use in UK beef evaluations. To undertake multitrait BLUP evaluations, it is necessary to have a full covariance matrix. This study outlines the approach taken to construct the full covariance matrices for the four beef breeds that most widely contribute to suckler beef cows in the United Kingdom. The maternal traits investigated were age at first calving, calving interval, lifespan, mature cow weight, 200-d weight, and calving difficulty. Three terminal sire traits (weight at 400 d, ultrasonic fat depth, and muscle score) were included to estimate covariances between the new and existing traits. A sire-maternal-grandsire model was used for the estimation procedure in a series of bivariate and multivariate models. A weighted bending procedure was employed to construct positive definite covariance matrices. Parameter estimates broadly agreed with literature values, although for some traits, literature information was very scarce. Some differences between parameters for different breeds were evident.     

Effects of lasalocid on circulating concentrations of leptin and insulin-like growth factor-I and reproductive performance of postpartum Brahman cows

Objectives were to determine effects of lasalocid on reproductive performance and serum concentrations of leptin and IGF-I, and to correlate concentrations of leptin and IGF-I with reproductive performance of beef cows. Forty-one purebred, multiparous Brahman cows were blocked to control (C; n = 20) or lasalocid (L; n = 21) treatments by BW, BCS, and predicted calving date. Treatment began 21 d before expected calving. Cows were each fed 1.4 kg daily of an 11:1 corn:soybean meal supplement, with the L group receiving 200 mg of lasalocid/cow daily. Cows and calves were weighed, and cow BCS was assessed at calving and at 28-d intervals thereafter. Blood samples were collected weekly precalving, at parturition, and twice weekly thereafter. Sterile marker bulls were maintained with cows for estrous detection. Six days after estrus, ovaries were evaluated for corpus luteum formation, and blood samples from d 6, 7, and 8 after estrus were collected. Serum samples were assayed for progesterone (P4), IGF-I, and leptin concentration. Progesterone concentrations > 1 ng/mL were considered indicative of a functional corpus luteum. Treatment ended after completion of a normal estrous cycle, and cows removed from treatment were placed with a fertile bull equipped with a chinball marker. There were no treatment differences in calving date, calf sex, cow BW, BCS, calf BW, calf ADG, or in serum concentrations of P4, IGF-I, or leptin. Prepartum cow ADG was increased (P < 0.01) in L cows and tended (P < 0.011) to be increased from calving to d 56 after calving in L cows. Postpartum interval (PPI) was not affected by treatment; however, a greater percentage (P < 0.05) of L cows conceived by 90 d after calving (43% L vs. 15% C). First-service conception rate tended (P < 0.08) to be greater in L vs. C cows (68 vs. 40%), but pregnancy rate was not different (P < 0.12; 86% for L vs. 65% for C). There were no treatment differences (P > 0.18) for serum IGF-I concentrations. At calving, leptin was positively correlated with IGF-I (P < 0.04; r = 0.32), BCS (P < 0.06; r = 0.29), and cow BW (P < 0.02; r = 0.36), and was negatively correlated with PPI (P < 0.06; r = -0.29). These results provide evidence that feeding an ionophore before calving and during the postpartum period may increase the number of cows that rebreed to maintain a yearly calving interval. Cows with higher concentrations of leptin postpartum may exhibit shorter PPI.     

黑龙江某规模化奶牛场荷斯坦牛群主要DHI指标分析

对黑龙江某规模化奶牛场2012年1—12月的DHI数据进行了系统的统计与分析,并将分析结果与国内外相关研究进展进行了对比。结果表明,该牛场的体细胞评分处于正常范围内,但存在泌乳高峰期到达时间晚、产犊间隔较长等问题。针对以上问题,提出了相应的改进措施。     

Some Factors Affecting Variation in Milk Yield in Crossbred Dairy Cows on Smallholder Farms in North-East Tanzania

A 2-year longitudinal survey was carried out to investigate factors affecting milk yield in crossbred cows on smallholder farms in and around an urban centre. Sixty farms were visited at approximately 2-week intervals and details of milk yield, body condition score (BCS) and heart girth measurements were collected. Fifteen farms were within the town (U), 23 farms were approximately 5 km from town (SU), and 22 farms approximately 10 km from town (PU). Sources of variation in milk yield were investigated using a general linear model by a stepwise forward selection and backward elimination approach to judge important independent variables. Factors considered for the first step of formulation of the model included location (PU, SU and U), calving season, BCS at calving, at 3 months postpartum and at 6 months postpartum, calving year, herd size category, source of labour (hired and family labour), calf rearing method (bucket and partial suckling) and parity number of the cow. Daily milk yield (including milk sucked by calves) was determined by calving year (p < 0.0001), calf rearing method (p = 0.044) and BCS at calving (p < 0.0001). Only BCS at calving contributed to variation in volume of milk sucked by the calf, lactation length and lactation milk yield. BCS at 3 months after calving was improved on farms where labour was hired (p = 0.041) and BCS change from calving to 6 months was more than twice as likely to be negative on U than SU and PU farms. It was concluded that milk production was predominantly associated with BCS at calving, lactation milk yield increasing quadratically from score 1 to 3. BCS at calving may provide a simple, single indicator of the nutritional status of a cow population.     

影响奶牛胎衣不下的主要因素分析

为了探讨影响奶牛胎衣不下的主要因素,笔者对新疆呼图壁种牛场产犊记录资料进行整理,通过SAS 8.1统计软件分析了产犊季节、胎次、性别、犊牛初生重等对奶牛胎衣不下的影响。结果表明,产犊季节对奶牛胎衣不下有极显著影响（P<0.01）,其中春季奶牛胎衣不下的发病率达到了40.72%;胎次对奶牛胎衣不下有极显著影响（P<0.01）,尤其是3胎以上的奶牛胎衣不下的发病率高于40%;犊牛性别和初生重对奶牛胎衣不下影响差异不显著（P>0.05）。     

Relationship between body condition score at calving and reproductive performance in young postpartum cows grazing native range

Body condition score is used as a management tool to predict competency of reproduction in beef cows. Therefore, a retrospective study was performed to evaluate association of BCS at calving with subsequent pregnancy rate, days to first postpartum ovulation, nutrient status (assessed by blood metabolites), and calf BW change in 2- and 3-yr-old cows (n = 351) managed and selected to fit their environment of grazing native range over 6 yr at the Corona Range and Livestock Research Center, Corona, NM. Cows were managed similarly before calving, without manipulation of management, to achieve predetermined BCS at parturition. Palpable BCS (scale of 1 to 9) were determined by 2 experienced technicians before calving. Cows were classified to 1 of 3 BCS groups prior calving: BCS 4 (mean BCS = 4.3 ± 0.02), 5 (mean BCS = 5.0 ± 0.03), or 6 (mean BCS = 5.8 ± 0.06). Cows were weighed weekly after calving and serum was collected once weekly (1 yr) or twice weekly (5 yr) for progesterone analysis to estimate first postpartum ovulation beginning 35 d postpartum. Year effects also were evaluated, with years identified as either above or below average precipitation. Days to first postpartum ovulation did not differ among calving BCS groups (P = 0.93). Pregnancy rates were not influenced by calving BCS (P = 0.83; 92%, 91%, 90% for BCS 4, 5, and 6, respectively). Days to BW nadir was not influenced by BCS at calving (P = 0.95). Cow BW was different at all measuring points (P < 0.01) with BCS 6 cows having the heaviest BW and cows with BCS 4 the lightest. Cows with calving BCS 4 and 5 lost more (P = 0.06) BW from the initiation of the study to the end of breeding than cows with BCS 6. However, cow BW change at all other measurement periods was not different (P ≥ 0.49) among calving BCS groups. Serum glucose and NEFA concentrations were not influenced by calving BCS (P ≥ 0.51). Calf BW at birth (P = 0.60), branding (55-d BW; P = 0.76), and weaning (205-d BW; P = 0.60) were not impacted by cow calving BCS. Body condition score did not influence overall pregnancy rates, indicating that young cows can have a reduced BCS and still be reproductively punctual. Therefore, these results indicate that reproductive performance of young cows with reduced BCS may not be affected when managed in extensive range conditions.