Heterophil/lymphocyte response of local Spanish breeds of laying hens to cold stress, heat stress, and water restriction.

Environmental challenges may adversely affect poultry production. Autochthonous breeds are particularly valuable in a climate change context because of adaptation to the local environment. The objective of the present study was to determine the impact of cold stress, water restriction, and heat stress on the stress response quantified by the heterophil to lymphocyte ratio (H/L) of ten local Spanish breeds of laying hens. Hens of these local breeds were consecutively subjected to three treatments: natural cold stress (2, 4, 6, 7, 9, and 13 °C), water restriction (with a duration of 2.5, 4.5, 7, 10, and 12 h, respectively), and natural heat stress (23, 26, 28, 30, 34, 38, 40, and 42 °C). During cold stress, H/L was higher at 9 and 13 °C than at 2, 4, and 6 °C, and higher at 9 °C than at 7 °C (P < 0.05). H/L values were similar during all water restriction levels. During heat stress, H/L was particularly elevated at temperatures higher than 40 °C (P < 0.05). Overall, Andaluza Azul, Andaluza Perdiz and Prat Codorniz showed lowest resilience to stress based on their H/L response, whereas Pardo de Leon, Villafranquina Roja, and Prat Leonada showed highest resilience.

Failing the four-gamete test enables exact phasing: the Corners' Algorithm.

BACKGROUND: Failing the four-gamete test for two polymorphic DNA markers is an indication that two or three rather than four haplotypes segregate in the population. The objective of this paper is to show that when just three haplotypes are segregating, all three haplotypes can be fully and unambiguously phase-resolved. THEORY AND METHODS: The Corners' Algorithm tests the four corners in a 3 × 3 table of two-locus genotypes. If one of the four corners is filled with zeroes, then the missing haplotype is identified and the phases of all three haplotypes can be unambiguously resolved for all individuals. Three applications of this method are proposed when the four-gamete test fails: (1) direct estimation of linkage disequilibrium (LD), (2) haplotype-based genome-wide association studies (GWAS) of three haplotypes (single-marker GWAS tests for two out of three haplotypes only), and (3) haplotyping of chromosomal regions that are comprised of pairs of single nucleotide polymorphisms (SNPs) that consist of just three haplotypes. An example based on 435 sows with performance records for total number of piglets born is used to illustrate the methods. RESULTS: Of 20,339 SNPs, approximately 50% of the pairs of flanking SNPs failed the four-gamete test. For those, the expectation maximization (EM) algorithm gave the same results. The average of the absolute value of the difference in r2 between flanking SNPs across the genome between the two methods was 0.00082. Single-marker GWAS (using two of three haplotypes) detected significant associations for total number of piglets born on chromosomes 1, 2, 6, 9, 10, 12, 13, 14, 15, and 18. Haplotype-based GWAS using the third haplotype resolved with the Corners' Algorithm detected additional significant associations for total number of piglets born on chromosomes 2, 5, 10, 13, 14, 15, and 18. Estimated substitution effects ranged from 0.40 to 1.35 piglets. Haplotyping of chromosomal regions that failed the four-gamete test for any pair of SNPs covered 961 Mb out of the 2249 Mb by the SNP array. CONCLUSIONS: The Corner's Algorithm allows to fully phase haplotypes when the four-gamete test fails. Longer haplotypes in chromosomal regions in which the four-gamete test fails for any pair of SNPs can be used as a multi-allelic marker with increased polymorphism information content.

Plasticity of feeding behaviour traits in response to production environment (temperate vs. tropical) in group-housed growing pigs.

Heat stress affects pig metabolism, health and welfare, resulting in reduced growth and important economic losses. The present experiment aimed to evaluate the effects of two climatic environments [temperate (TEMP) vs. tropical humid (TROP)] on feeding behaviour in growing pigs. The feeding behaviour traits were measured with automated feeders and included: daily feed intake, daily eating time, feeding rate, daily number of meals, feed intake per meal, and feeding time per meal. Pigs came from a backcross population between Large White (LW, heat sensitive) and Creole (CR, heat tolerant) pigs. The same 10 F1 LW × CR boars (sire families [SF]) were mated with related LW sows in each environment. Feeding behaviour was recorded for a total of 1,296 pigs (n = 634 pigs for TEMP and n = 662 pigs for TROP) between 11 and 23 weeks of age. Growth performance and thermoregulatory responses (rectal and skin temperatures) were also measured. Results show that TROP conditions affect feeding behaviour traits: animals had more meals per day but these meals were smaller both in duration and in size, resulting in lower daily feed intake and less time eating per day. Significant SF by environment (GxE) interactions were found for all feeding behaviour traits. When SF were distributed into robust and sensitive groups (previously defined according to performance and thermoregulatory traits), results showed group by environment interactions for all feeding traits, except meal frequency. Moreover, a significant difference in feeding rate between robust and sensitive group was detected in TEMP, suggesting that feeding rate may be a good candidate to evaluate heat tolerance.

Vector space algebra for scaling and centering relationship matrices under non-Hardy-Weinberg equilibrium conditions.

BACKGROUND: Scales are linear combinations of variables with coefficients that add up to zero and have a similar meaning to "contrast" in the analysis of variance. Scales are necessary in order to incorporate genomic information into relationship matrices for genomic selection. Statistical and biological parameterizations using scales under different assumptions have been proposed to construct alternative genomic relationship matrices. Except for the natural and orthogonal interactions approach (NOIA) method, current methods to construct relationship matrices assume Hardy-Weinberg equilibrium (HWE). The objective of this paper is to apply vector algebra to center and scale relationship matrices under non-HWE conditions, including orthogonalization by the Gram-Schmidt process. THEORY AND METHODS: Vector space algebra provides an evaluation of current orthogonality between additive and dominance vectors of additive and dominance scales for each marker. Three alternative methods to center and scale additive and dominance relationship matrices based on the Gram-Schmidt process (GSP-A, GSP-D, and GSP-N) are proposed. GSP-A removes additive-dominance co-variation by first fitting the additive and then the dominance scales. GSP-D fits scales in the opposite order. We show that GSP-A is algebraically the same as the NOIA model. GSP-N orthonormalizes the additive and dominance scales that result from GSP-A. An example with genotype information on 32,645 single nucleotide polymorphisms from 903 Large-White × Landrace crossbred pigs is used to construct existing and newly proposed additive and dominance relationship matrices. RESULTS: An exact test for departures from HWE showed that a majority of loci were not in HWE in crossbred pigs. All methods, except the one that assumes HWE, performed well to attain an average of diagonal elements equal to one and an average of off diagonal elements equal to zero. Variance component estimation for a recorded quantitative phenotype showed that orthogonal methods (NOIA, GSP-A, GSP-N) can adjust for the additive-dominance co-variation when estimating the additive genetic variance, whereas GSP-D does it when estimating dominance components. However, different methods to orthogonalize relationship matrices resulted in different proportions of additive and dominance components of variance. CONCLUSIONS: Vector space methodology can be applied to measure orthogonality between vectors of additive and dominance scales and to construct alternative orthogonal models such as GSP-A, GSP-D and an orthonormal model such as GSP-N. Under non-HWE conditions, GSP-A is algebraically the same as the previously developed NOIA model.

Impact of environmental temperature on production traits in pigs.

There is an urgent need to identify the effects of temperature on production traits. This study aimed to determine the impact of pig production in three environments (T°Cgrowing-°Cfattening-°Cfinishing = T24-24-21, T19-19-19, and T23-17-15) on growth curve parameters, body weight gain (DBWG), feed intake (DFI), and feed efficiency during the growing, fattening and finishing stages, and on carcass yield of primal cuts (ham, shoulder, and loin) in 158 Duroc × Iberian pigs. Maturation rate was higher in T23-17-15 than in T19-19-19 (P < 0.001). Pigs in T23-17-15 reached a lower mature body weight (P < 0.05). During the growing stage, pigs in T23-17-15 had higher DFI than those in T24-24-21 and T19-19-19 (P < 0.05); during the fattening stage, DFI was lowest in T24-24-21 (P < 0.001). In the growing stage, pigs had highest DBWG in the warmest environments (T24-24-21 and T23-17-15) and lowest in the coldest environment (T19-19-19; P < 0.001). Feed efficiency was highest in warmer environments (P < 0.01). Temperature T24-24-21 favored loin yield, T19-19-19 favored ham yield, and T23-17-15 favored shoulder yield (P < 0.01). The results imply a favorable effect of temperature on feed efficiency, however, possible negative implications for animal health and welfare should be considered.

Prospects for sustainability of pig production in relation to climate change and novel feed resources.

Pig production systems provide multiple benefits to humans. However, the global increase in meat consumption has profound consequences for our earth. This perspective describes two alternative scenarios for improving the sustainability of future pig production systems. The first scenario is a high input-high output system based on sustainable intensification, maximizing animal protein production efficiency on a limited land surface at the same time as minimizing environmental impacts. The second scenario is a reduced input-reduced output system based on selecting animals that are more robust to climate change and are better adapted to transform low quality feed (local feeds, feedstuff co-products, food waste) into meat. However, in contrast to the first scenario, the latter scenario results in reduced predicted yields, reduced production efficiency and possibly increased costs to the consumer. National evaluation of the availability of local feed and feedstuff co-product alternatives, determination of limits to feed sourced from international markets, available land for crop and livestock production, desired production levels, and a willingness to politically enforce policies through subsidies and/or penalties are some of the considerations to combine these two scenarios. Given future novel sustainable alternatives to livestock animal protein, it may become reasonable to move towards an added general premium price on 'protein from livestock animals' to the benefit of promoting higher incomes to farmers at the same time as covering the extra costs of, politically enforced, welfare of livestock animals in sustainable production systems. © 2020 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Effect of a dual enteric and respiratory pathogen challenge on swine growth, efficiency, carcass composition, and pork quality1.

The objective of this study was to determine the influence of a dual respiratory and enteric pathogen challenge on growth performance, carcass composition, and pork quality of high and low feed efficient pigs. Pigs divergently selected for low and high residual feed intake (RFI, ~68 kg) from the 11th generation of Iowa State University RFI project were used to represent high and low feed efficiency. To elicit a dual pathogen challenge, half of the pigs (n = 12/line) were inoculated with Mycoplasma hyopneumoniae (Mh) and Lawsonia intracellularis (MhLI) on days post-inoculation (dpi) 0. Pigs in a separate room of the barn were not inoculated and used as controls (n = 12/RFI line). Pigs were weighed and feed intake was recorded to calculate ADG, ADFI, and G:F for the acclimation period (period 1: dpi -21 to 0), during peak infection (period 2: dpi 0 to 42), and during the remaining growth period to reach market weight (period 3: dpi 42 to harvest). At ~125 kg, pigs were harvested using standard commercial procedures. Carcasses were evaluated for composition (weight, fat free lean, loin eye area, 10th rib fat depth) and meat quality (pH decline, temperature decline, Hunter L, a, and b, subjective color and marbling, star probe, drip loss, cook loss, proximate composition, and desmin degradation). Challenged pigs had lesser ADFI than controls during period 2 (P < 0.05), but had greater ADG and G:F during period 3 (P < 0.05). Selection for feed efficiency did not result in a differential response to MhLI (P > 0.05). Loin chops from the less feed efficient, high RFI pigs, had greater drip loss, greater cook loss, lesser moisture content, greater Hunter L values, and greater Hunter b values (P < 0.05) than loin chops from low RFI pigs. Infection status did not significantly affect carcass composition or pork quality traits (P > 0.05). These results indicate that a MhLI challenge early in growth did not significantly affect ultimate carcass composition or meat quality traits. Selection for greater feed efficiency in pigs did not affect their response to pathogenic challenge.

Extent of third-order linkage disequilibrium in a composite line of Iberian pigs.

BACKGROUND: Previous studies on linkage disequilibrium have investigated second order linkage disequilibrium in animal and plant populations. The objective of this paper was to investigate the genome-wide levels of third order linkage disequilibrium in a composite line founded by admixture of four Iberian pig strains. A model for the generation of third order linkage disequilibrium by population admixture is proposed. A computer Expectation-Maximization algorithm is developed and applied to the estimation of third order linkage disequilibrium at inter- and intra-chromosomal level using 26,347 SNPs typed in 306 sows. The relationship of third order linkage disequilibrium with physical distance was investigated over 35 million triplets in SSC12. Basic and normalized estimates of inter and intra-chromosomal third order linkage disequilibrium are reported. RESULTS: Genome-wide analyses revealed that third order linkage disequilibrium is rather common among linked loci in this Iberian pig line. It is shown that population admixture of multiple populations may explain the observed levels of third order linkage disequilibrium although it could be generated by genetic drift. Third order linkage disequilibrium decreases rapidly up to 4 Mb and then declines slowly. The short distances between consecutive markers explain the maintenance of the observed third order linkage disequilibria levels when using a model incorporating the break-up of disequilibrium by recombination. Genome-wide testing also revealed that only 3.6% of the normalized estimates were different from 1, - 1, 0, or from a not well-defined situation in which there is only one possible value for the third order linkage disequilibrium parameter, given allele frequencies and pairwise linkage disequilibria parameters. CONCLUSIONS: Third order linkage disequilibrium is common among linked markers in the analyzed pig line and may have been generated by population admixture of multiple populations or by genetic drift. As with second order linkage disequilibrium, the absolute value of the third order linkage disequilibrium decreases with physical distance. Normalization of third order linkage disequilibrium should be avoided for closely linked bi-allelic loci.

Impact of Mycoplasma hyopneumoniae and Lawsonia intracellularis on the performance of pigs divergently selected for feed efficiency.

Feed efficiency (FE) is a valuable trait, yet how genetic selection for enhanced FE affects other processes such as response to disease is unknown. Disease from endemic respiratory and enteric pathogens such as Mycoplasma hyopneumoniae (Mh) and Lawsonia intracellularis (LI) are common in swine production. Therefore, the aim of this study was to examine if pigs selected for high vs. low FE based on residual feed intake (RFI) respond differently to a dual respiratory and enteric challenge. Pigs selected for low RFI (LRFI, high FE) are considered more FE compared to their high RFI (HRFI, low FE) selected counterparts. Using a 2 × 2 factorial design, 25 littermate pairs from the HRFI and 25 littermate pairs from the LRFI line (barrows, 50 ± 7 kg BW) were selected, with one pig from each pair assigned to individual pens in either the challenge or the nonchallenge (control) rooms (n = 25 barrows/line/challenge). On days post inoculation (dpi) 0, the challenged pigs were inoculated with LI and Mh (MhLI). Feed intake, BW, fecal swabs, and serum samples were collected and recorded weekly for 42 d. On dpi -2 and 47, 14 littermate pairs (n = 7 barrows/line/challenge) were utilized for initial and final body composition scans using dual-energy X-ray absorptiometry to calculate longitudinal whole body tissue accretion rates for lean, protein, fat, and bone mineral content. Serum antibody levels and fecal shedding of LI were used to confirm infection. Control pigs remained negative by all measures during the 6-wk trial and MhLI inoculated pigs were confirmed positive via serological antibody responses by dpi 14 for LI and Mh. There were no interactions between RFI line and challenge status for any overall performance parameter (P > 0.05). The 6-wk MhLI challenge resulted in a 17% reduction in ADG, a 12% reduction in ADFI, and a 7% reduction in G:F vs. Controls (P < 0.05). In addition, compared to the Control pigs, MhLI challenge reduced lean, protein, and lipid accretion rates by 16% (P < 0.05). Genetic selection for high FE resulted in decreased ADFI and increased G:F (P < 0.01), but did not impact ADG or tissue accretion vs. low FE pigs. Collectively, these results demonstrate that a dual enteric and respiratory pathogen challenge reduced ADG, ADFI, G:F, and tissue accretion in growing pigs. Further, there was no evidence that selection for enhanced FE based on RFI index affects response to disease.

Effects of Diet and Genetics on Growth Performance of Pigs in Response to Repeated Exposure to Heat Stress.

Heat stress (HS) is one of the costliest issues in the U.S. pork industry. Aims of the present study were to determine the consequences of repeated exposure to HS on growth performance, and the effects of a high fiber diet, the genetic potential for high lean tissue accretion, and the genetic potential for residual feed intake (RFI) on resilience to HS. Barrows (n = 97) from three genetic lines (commercial, high RFI, low RFI) where subjected three times to a 4-day HS treatment (HS1, HS2, and HS3) which was preceded by a 9-day neutral (TN) adaptation period (TN1) and alternated by 7-day periods of neutral temperatures (TN2, TN3, and TN4). Body weight gain (BWG), feed intake (FI), feed conversion efficiency (FCE), RFI, and the drop in BWG and FI between TN and HS were estimated for each period, and slaughter traits were measured at the end of TN4. Commercial pigs had lower FI when fed a high fiber diet compared to a regular diet (2.70 ± 0.08 vs. 2.96 ± 0.08 kg/d; P < 0.05), while no differences were found for BWG, RFI or FCE. HS reduced FI, BWG, and FCE, increased RFI, and resulted in leaner pigs that generate smaller carcasses at slaughter. In TN, commercial pigs grew faster than the low and high RFI pigs (1.22 ± 0.06 vs. 0.720 ± 0.05 and 0.657 ± 0.07; P < 0.001) but growth rates were not significantly different between the lines during HS. Growth rates for the low RFI and high RFI pigs were similar both during TN and during HS. Pigs of interest for genetic improvement are those that are able to maintain growth rates during HS. Our results show that response in growth to HS was repeatable over subsequent 4-d HS cycles, which suggests the potential for including this response in the breeding index. The best performing animals during HS are likely those that are not highly superior for growth in TN.

A Hypothesis and Review of the Relationship between Selection for Improved Production Efficiency, Coping Behavior, and Domestication.

Coping styles in response to stressors have been described both in humans and in other animal species. Because coping styles are directly related to individual fitness they are part of the life history strategy. Behavioral styles trade off with other life-history traits through the acquisition and allocation of resources. Domestication and subsequent artificial selection for production traits specifically focused on selection of individuals with energy sparing mechanisms for non-production traits. Domestication resulted in animals with low levels of aggression and activity, and a low hypothalamic-pituitary-adrenal (HPA) axis reactivity. In the present work, we propose that, vice versa, selection for improved production efficiency may to some extent continue to favor docile domesticated phenotypes. It is hypothesized that both domestication and selection for improved production efficiency may result in the selection of reactive style animals. Both domesticated and reactive style animals are characterized by low levels of aggression and activity, and increased serotonin neurotransmitter levels. However, whereas domestication quite consistently results in a decrease in the functional state of the HPA axis, the reactive coping style is often found to be dominated by a high HPA response. This may suggest that fearfulness and coping behavior are two independent underlying dimensions to the coping response. Although it is generally proposed that animal welfare improves with selection for calmer animals that are less fearful and reactive to novelty, animals bred to be less sensitive with fewer desires may be undesirable from an ethical point of view.

Allometric scaling of the elevation of maternal energy intake during lactation.

BACKGROUND: In most mammals, lactating mothers dramatically increase their food intake after parturition and reach a peak intake rate after a certain time while their offspring continue to grow. A common view, perpetuated by the metabolic theory of ecology, is that the allometric scaling of maternal metabolic rate with body mass limits the changes in energy intake and expenditure. Therefore these potential effects of metabolic scaling should be reflected in the elevation of maternal energy intake during lactation. To test this hypothesis, we collected published data on 24 species (13 domesticated) and established scaling relationships for several characteristics of the patterns of energy intake elevation (amplitude of the elevation, time to peak, and cumulative elevation to peak). RESULTS: A curvilinear allometric scaling relationship with maternal body mass (in double-logarithmic space) was found for the amplitude of maternal energy intake elevation, similarly to what has been observed for scaling relationships of basal metabolic rate in non-breeding mammals. This result indirectly supports the metabolic theory of ecology. However, this curvilinear allometric scaling does not seem to drive the scaling relationships found for the other characteristics of maternal energy intake. Both the duration and shape of the energy intake patterns showed substantial variation independently of species' body mass. CONCLUSIONS: Data available for a few mammals, mostly domesticated, provides little evidence for the hypothesis that a single law of metabolic scaling governs the elevation of maternal energy intake after parturition. Obtaining further food intake data in wild species will be crucial to unravel the general mechanisms underlying variation in this unique adaptation of mammalian females.

Genotype by environment interaction and breeding for robustness in livestock.

The increasing size of the human population is projected to result in an increase in meat consumption. However, at the same time, the dominant position of meat as the center of meals is on the decline. Modern objections to the consumption of meat include public concerns with animal welfare in livestock production systems. Animal breeding practices have become part of the debate since it became recognized that animals in a population that have been selected for high production efficiency are more at risk for behavioral, physiological and immunological problems. As a solution, animal breeding practices need to include selection for robustness traits, which can be implemented through the use of reaction norms analysis, or though the direct inclusion of robustness traits in the breeding objective and in the selection index. This review gives an overview of genotype × environment interactions (the influence of the environment, reaction norms, phenotypic plasticity, canalization, and genetic homeostasis), reaction norms analysis in livestock production, options for selection for increased levels of production and against environmental sensitivity, and direct inclusion of robustness traits in the selection index. Ethical considerations of breeding for improved animal welfare are discussed. The discussion on animal breeding practices has been initiated and is very alive today. This positive trend is part of the sustainable food production movement that aims at feeding 9.15 billion people not just in the near future but also beyond.

Haplotype phasing after joint estimation of recombination and linkage disequilibrium in breeding populations.

A novel method for haplotype phasing in families after joint estimation of recombination fraction and linkage disequilibrium is developed. Results from Monte Carlo computer simulations show that the newly developed E.M. algorithm is accurate if true recombination fraction is 0 even for single families of relatively small sizes. Estimates of recombination fraction and linkage disequilibrium were 0.00 (SD 0.00) and 0.19 (SD 0.03) for simulated recombination fraction and linkage disequilibrium of 0.00 and 0.20, respectively. A genome fragmentation phasing strategy was developed and used for phasing haplotypes in a sire and 36 progeny using the 50 k Illumina BeadChip by: a) estimation of the recombination fraction and LD in consecutive SNPs using family information, b) linkage analyses between fragments, c) phasing of haplotypes in parents and progeny and in following generations. Homozygous SNPs in progeny allowed determination of paternal fragment inheritance, and deduction of SNP sequence information of haplotypes from dams. The strategy also allowed detection of genotyping errors. A total of 613 recombination events were detected after linkage analysis was carried out between fragments. Hot and cold spots were identified at the individual (sire level). SNPs for which the sire and calf were heterozygotes became informative (over 90%) after the phasing of haplotypes. Average of regions of identity between half-sibs when comparing its maternal inherited haplotypes (with at least 20 SNP) in common was 0.11 with a maximum of 0.29 and a minimum of 0.05. A Monte-Carlo simulation of BTA1 with the same linkage disequilibrium structure and genetic linkage as the cattle family yielded a 99.98 and 99.94% of correct phases for informative SNPs in sire and calves, respectively.

Hematologic and IgG responses of heifers experimentally infected with the agent of epizootic bovine abortion.

BACKGROUND: Epizootic bovine abortion (EBA) is a tick-transmitted abortive disease of beef cattle in the western United States. Infected cattle do not have clinical signs until abortion occurs, usually within the last trimester of gestation. There is little information on the hematologic response of the dam following infection. OBJECTIVE: The purpose of this study was to determine if changes in blood leukocytes and serum IgG concentrations could be detected following experimental infection of pregnant heifers with the etiologic agent of EBA (aoEBA). METHODS: Twelve Angus heifers were infected during gestation with the aoEBA using an inoculum prepared from the thymus of an infected fetus. Five pregnant heifer controls were given an inoculum prepared from the thymus of an aoEBA-negative calf. PCVs, total and differential leukocyte counts, and serum IgG concentrations were measured weekly following administration of the inocula until abortion or calving. Gross and microscopic examinations were performed on all aborted fetuses to confirm infection. RESULTS: Eleven of 12 heifers in the treatment group aborted, and significant findings were decreased lymphocyte counts at weeks 1 and 14 postinoculation and increased monocyte counts at week 4 compared with control animals. Serum IgG concentrations were significantly increased at weeks 6-8 and 11 in the treatment group. CONCLUSIONS: Leukogram changes are subtle in infected cattle. Future research efforts should be aimed at development of an antibody test specific for detection of previously infected animals, which could graze safely on EBA-endemic pastures.

Immune response from a resource allocation perspective.

The immune system is a life history trait that can be expected to trade off against other life history traits. Whether or not a trait is considered to be a life history trait has consequences for the expectation on how it responds to natural selection and evolution; in addition, it may have consequences for the outcome of artificial selection when it is included in the breeding objective. The immune system involved in pathogen resistance comprises multiple mechanisms that define a host's defensive capacity. Immune resistance involves employing mechanisms that either prevent pathogens from invading or eliminate the pathogens when they do invade. On the other hand, tolerance involves limiting the damage that is caused by the infection. Both tolerance and resistance traits require (re)allocation of resources and carry physiological costs. Examples of trade-offs between immune function and growth, reproduction and stress response are provided in this review, in addition to consequences of selection for increased production on immune function and vice versa. Reaction norms are used to deal with questions of immune resistance vs. tolerance to pathogens that relate host health to infection intensity. In essence, selection for immune tolerance in livestock is a particular case of selection for animal robustness. Since breeding goals that include robustness traits are required in the implementation of more sustainable agricultural production systems, it is of interest to investigate whether immune tolerance is a robustness trait that is positively correlated with overall animal robustness. Considerably more research is needed to estimate the shapes of the cost functions of different immune strategies, and investigate trade-offs and cross-over benefits of selection for disease resistance and/or disease tolerance in livestock production.

Food resource allocation patterns in lactating females in a long-term selection experiment for litter size in mice.

Resource allocation patterns, as quantified by residual food intake (RFI), and the consequences for offspring development were investigated during lactation in 96 females of a mouse line selected for 104 generations for high litter size at birth (S-line) and in 87 females of a non-selected control line (C-line). Litters of 45 C-line dams (Cs) and 48 S-line dams (Ss) were standardised (s) at birth; other dams (ns) supported total number of pups born (Cns and Sns, respectively). RFI during lactation was significantly lower in Sns-dams than in C-line dams and Sns-dams. After weaning Sns-dams seemed to be able to restore the negative resource situation. Sns-pups were about 25% less mature than Cns-pups at all times. Maturity was similar for Cs- and Ss-pups from 2 d in lactation on, and about 18% and 53% higher than Cns- and Sns-pups. The pre-weaning mortality rate was significantly higher in Sns-litters (35.6 +/-2.76) than in Cns-litters (4.95 +/-2.23). The results suggest that S-line dams allocated considerably more resources to maintenance of offspring than C-line dams. This was insufficient to provide the offspring with an adequate amount of resources, resulting in reduced pup development and increased pre-weaning mortality rates.