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Selection for sow longevity.

Sow longevity plays an important role in economically efficient piglet production because sow longevity is related to the number of piglets produced during its productive lifetime; however, selection for sow longevity is not commonly practiced in any pig breeding program. There is relatively little scientific literature concerning the genetic parameters (genetic variation and genetic correlations) or methods available for breeding value estimation for effective selection for sow longevity. This paper summarizes the current knowledge about the genetics of sow longevity and discusses the available breeding value estimation methods for sow longevity traits. The studies in the literature clearly indicate that sow longevity is a complex trait, and even the definition of sow longevity is variable depending on the researcher and research objective. In general, the measures and analyses of sow longevity can be divided into 1) continuous traits (e.g., productive lifetime) analyzed with proportional hazard models; and 2) more simple binary traits such as stayability until some predetermined fixed parity. Most studies have concluded that sufficient genetic variation exists for effective selection on sow longevity, and heritability estimates have ranged between 0.02 and 0.25. Moreover, sow longevity has shown to be genetically associated with prolificacy and leg conformation traits. Variable results from previous research have led to a lack of consensus among swine breeders concerning the valid methodology of estimating breeding values for longevity traits. One can not deny the superiority of survival analysis in the modeling approach of longevity data; however, multiple-trait analyses are not possible using currently available survival analysis software. Less sophisticated approaches have the advantage of evaluating multiple traits simultaneously, and thus, can use the genetic associations between sow longevity and other traits. Additional research is needed to identify the most efficient selection methods for sow longevity. Future research needs to concentrate on multiple trait analysis of sow longevity traits. Moreover, because longevity is a fitness trait, the nonadditive genetic effects (e.g., dominance) may play important role in the inheritance of sow longevity. Currently, not a single estimate for dominance variance of sow longevity could be identified from the scientific literature.

Animal Husbandry↗

Longevity and efficiency associated with age structures of female pigs and herd management in commercial breeding herds.

Annual performance measurements, age structures of female pig inventories, and by-parity culling rates were abstracted from data files of 110 herds that participated in a data-share program in Japan. Parity at culling was used as a prime measurement of longevity, whereas pigs weaned x mated female(-1) x year(-1) (PWMFY) was a prime measurement of reproductive efficiency. High or low longevity herds were based on the greatest 50% of the herds or the remaining herds ranked by parity at culling, whereas high or low reproductive efficiency herds were grouped according to the greatest 50% of the herds or the remaining herds ranked by PWMFY. Measurements were analyzed as a 2 x 2 factorial arrangement, using the main effects of the 2 herd groups of longevity (high or low) and reproductive efficiency (high or low). Means of parity at culling and PWMFY were 4.6 (SD = 0.82) and 21.2 (SD = 3.02), respectively. The high longevity group had 1.27 greater parities at culling than the low longevity group (P < 0.05), but no differences between the high and low longevity groups were found in PWMFY (P = 0.21). No differences between the high and low efficiency groups were found in parity at culling (P = 0.50). No interactions between the longevity and efficiency groups were found on any longevity or efficiency measurement (P > 0.20). In herd management, the percentage of reserviced females and the percentage of multiple matings were associated with the longevity group and the efficiency group (P < 0.05). The high longevity group had lower culling rates in parity 0 to 6 than the low longevity group (P < 0.05), whereas no differences between the low and high efficiency groups were found in culling rates in parity 0 to 2 (P > 0.20). This study suggests that measures to achieve longevity and high reproductive efficiency in breeding herds do not conflict and that high reproductive efficiency and high longevity can be achieved.

Aging↗

Genetic parameters of longevity traits of an upgrading population of dairy cattle.

Longevity reflects the ability of a cow to avoid being culled for low production, low fertility, or illness. Longevity could be used in breeding programs if genetic parameters were known. Various measures are used for longevity. In this study, lifetime measures including number of lactations, total milk production, number of days in lactation, herd life, and length of productive life were analyzed. Also analyzed were stayability measures (dead or alive) to 36, 48, 60, or 72 mo of age and to 12, 24, 36, or 48 mo of productive life. Measures of longevity were also analyzed after correction for milk production during first lactation (functional longevity traits). Data on 1,72,988 cows were used to calculate means for longevity traits per year of birth. All cows were known to have been culled. Longevity decreased from 1978 through 1984 and increased in 1985. Possible causes for the decrease of longevity were implementation of the quota system and introduction of Holstein genes. heritabilities of longevity traits were estimated for cows born in 1985 (38,957 records), 1982 (166,324 records), and 1978 (94,935 records) after data were edited to require at least 25 daughters per sire and 10 cows per herd. Phenotypic and genetic correlations were estimated for the 1985 data. Heritability estimates differed between years of birth, and estimates of functional traits were lower than those of uncorrected longevity traits. Genetic correlations between uncorrected longevity traits were high (0.733 to 1.000); phenotypic correlations were lower (0.131 to 0.980). Genetic correlations between uncorrected and functional longevity traits were high (0.577 to 0.975).

Aging↗

Genetic evaluation of length of productive life including predicted longevity of live cows.

Complete longevity data are available too late for most sire selection. Earlier selection is possible using correlated traits, nonlinear evaluation of censored data, or predicted longevities for live cows in addition to completed longevity data. Completed longevity was defined as total months in milk by 84 mo of age. Predicted longevity was computed by multiple regression from cows alive at six different ages. Variables included age at first calving, standardized first lactation milk yield (optional), lactation status (dry or milking), current months in milk, current months dry, and cumulative months in milk. Completed longevity data for dead cows were then merged with predicted longevity data for live cows. A total of 1,984,038 Holstein cows born from 1979 to 1983 were included and represented 1911 sires, each with at least 70 daughters. Heritability of longevity increased gradually from .03 at 36 mo to .08 at 84 mo. Phenotypic correlations of early with completed longevity ranged from .59 to .99; genetic correlations ranged from .92 to 1.00. Inclusion of yield for projection increased heritability at 36 mo substantially but decreased genetic correlation with complete longevity information. Expansion and weighting factors will allow predicted records to be used in longevity evaluations, which is similar to the procedures that allow predicted 305-d yields to be included in yield evaluations.

Aging↗

Quantitative trait loci analysis of leaf and plant longevity in Arabidopsis thaliana.

The natural variation in leaf and plant longevity in Arabidopsis thaliana was analysed in a set of 45 ecotypes and 155 recombinant inbred lines derived from a Cape Verde Islands (Cvi) x Landsberg erecta (Ler) cross. Post-bolting longevity was inversely related to time to flowering and rosette leaf number in the set of 45 ecotypes, with Cvi having the longest and Ler the shortest post-bolting longevity. The recombinant inbred line population was tested under low or high soil nutrient levels (LN or HN, respectively). Three quantitative trait loci (QTL), one in chromosome 3 and two in chromosomes 1 and 5, were associated with longevity of the 6th rosette leaf under LN and HN, respectively. Four QTL for post-bolting longevity were found in chromosomes 1, 3, 4, and 5, and two in chromosomes 1 and 5 under LN and HN, respectively. An epistatic interaction affecting post-bolting longevity under LN, but not HN, was detected. Ler and Cvi carry a mix of increasing and decreasing alleles for the QTL affecting longevity of the 6th leaf and post-bolting longevity. Longevity of the 6th rosette leaf was associated with different QTL than post-bolting longevity, and it was affected by different QTL depending on nutrient availability. By contrast, the major QTL affecting post-bolting longevity exerted significant effects irrespective of soil nutrient availability.

Arabidopsis↗

Canopy dynamics and aboveground production of five tree species with different leaf longevities.

Canopy dynamics and aboveground net primary production (ANPP) were studied in replicated monospecific and dual-species plantations comprised of species with different leaf longevities. In the monospecific plantations, leaf longevity averaged 5, 6, 36, 46 and 66 months for Quercus rubra L., Larix decidua Miller, Pinus strobus L., Pinus resinosa Ait. and Picea abies (L.) Karst., respectively. Specific leaf area, maximum net photosynthesis per unit mass (A/mass), leaf N per unit mass (N(leaf)/mass) and maximum net photosynthesis on a leaf N basis (A/N(leaf)) were inversely correlated to leaf longevity (r(2) = 0.92-0.97, 0.91, 0.88 and 0.80, respectively). Maximum net photosynthesis per unit area (A/area) was not correlated to leaf longevity, whereas leaf N per unit area (N(leaf)/area) was positively correlated to leaf longevity (r(2) = 0.95). For a similar-diameter conifer, species with long-lived foliage supported a greater foliage mass than species with short-lived foliage; however, Quercus rubra did not follow this pattern. At the stand level, total foliage mass ranged from 3.3 to 30.5 Mg ha(-1) and was positively correlated (r(2) = 0.97) to leaf longevity. Leaf area index (LAI) was also positively correlated (r(2) = 0.82) to leaf longevity. Production efficiency (ANPP/LAI) was inversely related to leaf longevity and positively related to A/mass. Aboveground biomass and net primary production differed significantly (P < 0.05) among the five species but were not correlated to leaf longevity, total foliage mass or leaf area. In monospecific plantations, stem NPP for Larix decidua was 17% greater than for Pinus strobus and 14% less than for Picea abies, but in mixed-species plantations stem NPP for Larix decidua was 62 and 85% greater than for Pinus strobus and Picea abies, respectively. Similar aboveground net primary production rates can be attained by tree species with different leaf longevities because of trade-offs resulting from different structural and physiological leaf and canopy characteristics that are correlated to each other and to leaf longevity.

Journal Article↗

Leaf longevity of western red cedar (Thuja plicata) increases with depth in the canopy.

Leaf longevity determines the annual cost of replacing foliage biomass and influences water and nutrient budgets. Longevity is readily estimated in most evergreen species by annual bud scars along the shoot. However, some species with indeterminate growth do not show these annual markers, making estimation of longevity difficult. One of these species is the widespread and economically valuable western red cedar (Thuja plicata J. Donn ex D. Don), for which no dependable estimates of leaf longevity exist. In this study, we estimated leaf longevity for western red cedar by counting growth rings in shoots at the point of leaf abscission. Estimates were obtained on 26 dominant or codominant trees growing in natural stands in a montane forest in northern Idaho, USA. Leaf longevity averaged 8.9 (SE = 0.2) years, but it strongly increased with depth in the canopy (0.3 year m(-1); mean crown depth was 15 m), increasing from a mean of 6.8 years in the upper third of the canopy to 10.6 years in the lower third. The increase in longevity with depth in the crown is consistent with many reports showing that longevity increases in resource-limiting environments. Longevity did not vary significantly with altitude or solar insolation in these montane forests. Among stand-level variables, longevity was correlated only with leaf area index: it increased slightly in stands with high leaf area indices. This approach to longevity estimation may be useful for any species that produces annual rings but no obvious bud scars, including many Cupressaceae species.

Altitude↗

Longevity network: construction and implications.

The vast majority of studies on longevity have focused on individual genes/proteins, without adequately addressing the possible role of interactions between them. This study is the first attempt towards constructing a "longevity network" via analysis of human protein-protein interactions (PPIs). For this purpose, we (i) compiled a complete list of established longevity genes from different species, including those that most probably affect the longevity in humans, (ii) defined the human orthologs of the longevity genes, and (iii) determined whether the encoded proteins could be organized as a network. The longevity gene-encoded proteins together with their interacting proteins form a continuous network, which fits the criteria for a scale-free network with an extremely high contribution of hubs to the network connectivity. Most of them have never been annotated before in connection with longevity. Remarkably, almost all of the hubs of the "longevity network" were reported to be involved in at least one age-related disease (ARD), with many being involved in several ARDs. This may be one of the ways by which the proteins with multiple interactions affect the longevity. The hubs offer the potential of being primary targets for longevity-promoting interventions.

Aging↗

Beneficial influences of systemic cooperation and sociological behavior on longevity.

During his long research career in the field of aging, Dr Bernard Strehler developed a series of theories concerning the identity of genes that can promote longevity and their role in natural selection. As a tribute to Dr Strehler, we have taken this opportunity to summarize a selection of these theories and to illustrate how these insights have influenced our search for longevity genes within the immune system. The identification of longevity genes has proven difficult. We believe that, at least in part, this reflects the emphasis on the concept of survival of the 'physically' fittest. We have used the immune system as a model to demonstrate that, over and above the self-evident advantage of those genes that contribute the attributes commonly associated with survival of the 'physically' fittest, those genes that lead to a predisposition to cooperate also confer a competitive survival advantage. As the acquisition of cooperativity in a society is linked to support mechanisms provided by older individuals, the search for longevity genes should not be limited to those genes that are associated with extended expression of a youthful phenotype. Rather these studies should be expanded to include identification of those genes that regulate physiologic parameters that affect individual longevity, even if they do not correspond with the traditional view of reproductive competitiveness. At the societal level, longevity genes may encode attributes that regulate sociologic or psychological parameters that may contribute to a tendency to non-aggressive or cooperative behavior that leads to achievement of common goals necessary for the survival of the species. This view of the selection for longevity impacts the analysis of longevity genes and aging at the organismal level. Dr Strehler viewed organismal aging as an integrated functional state, in which he conceived the outcome as reflecting the net balance of functional decrementers and evolved compensatory features. We propose that, in more evolved species, the longevity genes will be those genes, or sets of genes, that counterbalance of age-related functional decrementers with the age-related manifestation of evolved compensatory features. Thus, as illustrated here through analysis of the immune system, the longevity genes may well be those genes that promote overall systemic cooperation and compensation within the immune system and associated systems, rather than the genes that prevent age-related alterations in only one or a limited number of pathways.

Aging↗

Variation in needle longevity of Pinus tabulaeformis forests at different geographic scales.

Needle longevity of conifer species is known to increase with latitude, but little is known about intraspecific variation and associated factors within a location. Chinese pine (Pinus tabulaeformis Carr.) forests were investigated to identify patterns of needle longevity at seven sites with distinctive climatic conditions along a latitudinal gradient (33-38 degrees N) in Shaanxi province, northwest China. A demographic approach was used to quantify needle longevity as an index of entire foliage population adjusted to needle-age-specific mortality rates. There were significant differences in needle longevity of Chinese pine stands across sites and across sample plots within a site. Individual tree needle longevity ranged from 0.62 to 3.75 years for 276 samples across sites. Needle longevity increased with latitude (R2 = 0.40, P < 0.0001), but decreased with mean January temperature (R2 = 0.63, P < 0.0001). Foliage retention of Chinese pine stands at the regional level was generally associated with climatic variability, indicating that variation in needle longevity was primarily an environmental acclimation to low temperature in winter. Stand characteristics were closely associated with needle longevity at three sites located within the same climatic zone. Needle longevity was positively correlated with tree age (R2 = 0.48, P < 0.0001) and stand density (R2 = 0.26, P = 0.0015) at Huanglong and Huangling, respectively, whereas it was negatively associated with total tree height at Zhidan (R2 = 0.50, P < 0.0001). It is concluded that, at the stand and individual tree level, intraspecific variation in needle longevity is most likely a result of adaptation to patchy microsite environments.

China↗

Regulation of longevity and stress resistance: a molecular strategy conserved from yeast to humans?

Recent studies implicate similar proteins in the regulation of longevity in organisms ranging from yeast to mice. Studies in yeast and worms suggest that inactivation of glucose or insulin/insulin-like growth factor-l (IGF-1) signaling pathways extends longevity by causing a shift from a reproductive phase to a non-reproductive maintenance phase involving the expression of many genes. These stress resistance pathways appear to have evolved to induce maintenance systems and promote longevity during periods of starvation. In yeast, mutations that decrease the activity of glucose signaling pathways extend longevity by activating stress resistance transcription factors that regulate the expression of genes involved in antioxidant and heat protection, glycogen storage, protein degradation, DNA repair, and metabolism. A remarkably similar set of proteins regulated by growth factors that control glucose metabolism is implicated in life span extension in worms, and possibly in flies and mice. Studies in worms and flies point to secondary hormones as mediators of the effect of insulin/ IGF-1 signaling on longevity, whereas studies in yeast and mammalian cells indicate that glucose or insulin/ IGF-1 may decrease longevity by directly down-regulating stress resistance genes. In yeast, longevity mutations postpone superoxide toxicity and mitochondrial damage. However, the small life span extension caused by the overexpression of superoxide dismutases and catalase in yeast and flies indicates that increased antioxidant protection alone cannot be responsible for the major life span extension caused by signal transduction mutations. Although we are only beginning to understand the molecular mechanisms that mediate life span extension, the similarities between longevity regulatory pathways in organisms ranging from yeast to mice suggest that insulin/ IGF-1 signaling pathways may also regulate cell damage and longevity in humans.

Animals↗

Heat-induced hormesis in longevity of two sibling Drosophila species.

Previous work showed that mild-heat stress induces longevity hormesis in a model organism, D. melanogaster. Here we compared the possible heat-induced hormesis in longevity of other species of Drosophila, D. buzzatii and its sibling species D. koepferae, in a single-sex environment. Possible correlations between longevity and heat-stress resistance were also tested by measuring longevity, heat-knockdown resistance and the heat-induced Hsp70 expression for each species in a common environment. D. buzzatii was longer lived than D. koepferae at benign temperature. Knockdown resistance to heat stress was positively correlated to longevity within species. However, the shorter-lived species was more resistant to knockdown by heat stress than the longer-lived species. The heat-induced Hsp70 expression was similar between species. A heat-shock treatment (37 degrees C for 1 h at 4 days of age) extended mean longevity in the longer lived species but not in the shorter lived species. In D. koepferae, the demographic rate of senescence decreased but the baseline mortality rate increased by heat-shock, resulting in no extension of mean longevity. Sympatric populations of closely related species can be differentially sensitive to temperature and exhibit different patterns of 37 degrees C-induced hormesis in demographic senescence and longevity. The results also show that positive correlations between stress resistance and life span within species can shift in sign across closely related species. Finally, this study shows that heat-induced hormesis in longevity can be found across different Drosophila species, as hormetic effects are not limited to the previously studied D. melanogaster.

Aging↗

Interactive environmental and genetic effects on longevity in the male rat: litter size, exercise, electric shocks and castration.

The effects on longevity of manipulating litter size during rearing and of postweaning forced exercise were observed in intact and castrated male rats. Random-bred sib quintets of newborn littermates were reared in reduced litters or normal litters (5 or 6, or 10-12 young, respectively). The quintets were split into five postweaning treatments: untreated (control), exercised (forced to run by electric shocks), shocked (without exercise), castrated, and castrated and exercised. Reduced litter size increased weaning weight from 46 to 56 g and diminished mean longevity from 724 to 620 days. The decrease in longevity due to reduced litter size was greater in the control (204 days) than in the other postweaning treatments (42 to 142 days). The postweaning treatments affected longevity only in the reduced-litter males; in these, exercise and mere shocks appeared to increase longevity. The variation in longevity among quintets was large and the expression of genetic longevity interacted with the treatments. The regression of individual on mean sib longevity was .89 in the castrates but only .10 in the controls; in exercised and shocked males it was .58 in normal-litter but only .09 in reduced-litter sibs indicating an interaction between pre- and postweaning treatments. The implications of the results on the interpretation of longevity studies are discussed.

Adipose Tissue↗

Is melatonin circadian rhythm a physiological feature associated with healthy longevity? A study of long-living subjects and their progeny.

The study investigates the circadian rhythm (CR) of urinary 6-sulphatoxy-melatonin (aMT6s) in long-living (longevous) subjects and their progeny. The aim is to detect whether or not the melatonin CR is a physiological feature associated with healthy longevity. The aMT6s CR was investigated in 10 longevous subjects, 8 of their children and 9 of their grandchildren, all in good health. Control data were obtained respectively from 13 adult subjects and 9 young subjects, in good health, but characterized by a negative family history for longevity. All the subjects were born and living in the same city. The study was performed in the summer of 1996. The aMT6s CR was found to persist in longevous subjects, being characterized by a lower mesor and amplitude. The aMT6s CR was found not to show properties consistently different in children and grandchildren as compared respectively to their adult and young controls. Because of its preservation in longevous subjects, it can be argued that the melatonin CR is a physiological feature associated with healthy longevity. Because of the comparability of aMT6s CR in children and grandchildren, with respect to their controls without a positive family history of longevity, it can be argued that the melatonin CR is not a marker that can be used for an earlier identification of the candidates for longevity.

Adult↗