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PubMed2005

Concurrent natural and sexual selection in wild male sockeye salmon, Oncorhynchus nerka.

Concurrent natural and sexual selection have been inferred from laboratory and comparative studies in a number of taxa, but are rarely measured in natural populations. Because the interaction of these two general categories of selection may be complex when they occur simultaneously, empirical evidence from natural populations would help us to understand this interaction and probably give us greater insight into each separate episode as well. In male sockeye salmon, sexual selection for larger body size has been indicated in both deep and shallow water habitats. However, in shallow habitats male sockeye are generally smaller and less deep-bodied than in deep habitats, a difference that has been ascribed to natural selection. We measured concurrent natural and sexual selection in two years on breeding male sockeye salmon with respect to body size, body shape, and time of arrival to the breeding grounds. Natural selection was variable in effect and sexual selection was variable in intensity in these two years. The patterns of selection also appear to be interdependent; areas where predation on spawning adults is not intense have yielded different patterns of sexual selection than those measured here. It appears that some of the body shape differences in sockeye salmon associated with different spawning habitats, which were previously attributed to selective mortality, may be a result of different patterns of sexual selection in the different habitats. Total selection resulting from the combination of both natural and sexual selection was less intense than either natural or sexual selection in most cases. Measurement of concurrent selection episodes in nature may help us to understand whether the pattern of differential sexual selection is common, and whether observed patterns of habitat-related differentiation may be due to differences in sexual selection.

Alaska↗
PubMed1993

Selective association of lipoprotein cholesteryl esters with liver plasma membranes.

High-density lipoprotein (HDL) cholesteryl esters are taken up by hepatocytes without parallel uptake of HDL apolipoproteins. This selective uptake of HDL cholesteryl esters is mediated by a non-endocytotic mechanism. Recently, selective uptake of cholesteryl esters also from low-density lipoprotein (LDL) was demonstrated. In this study, the role of the plasma membrane in selective uptake by the liver was investigated. Plasma membranes were prepared from rat liver or from human Hep G2 hepatoma cells. Human HDL3 (d = 1.125-1.21 g/ml) was either radioiodinated or labeled with [3H]cholesteryl oleate. Human low-density lipoprotein (d = 1.019-1.05 g/ml) was labeled in its protein and in its lipid moiety as well. Labeled lipoproteins, unlabeled lipoproteins and membranes were incubated. After separation by ultracentrifugation, apparent lipoprotein particle association with membranes was determined. Plasma membranes from rat liver and Hep G2 cells bound 125I-HDL3, indicating specific HDL3 particle binding. With both types of membrane, apparent HDL3 particle association according to [3H]cholesteryl oleate-labeled HDL3 was in significant excess on that due to 125I-HDL3. This indicates selective, i.e., particle binding independent, association of cholesteryl esters with the membrane. Excess unlabeled HDL3 competed for selective association, indicating a specific process. Selective association of HDL3 cholesteryl esters was concentration-, time-, temperature-dependent; however, parameters differed from HDL3 particle binding. HDL3 was modified by nitration; this modification inhibited HDL3 particle binding in contrast to unchanged selective association. These results suggested distinct membrane sites for HDL3 particle binding and selective cholesteryl ester association. Regulation of selective association was investigated. Hep G2 cells were cholesterol-loaded or cholesterol-depleted. Cellular cholesterol-loading down-regulated selective association of HDL3 cholesteryl esters with isolated membranes prepared from these cells. In parallel, selective uptake of HDL3 cholesteryl esters by Hep G2 cells was down-regulated in cholesterol-loaded cells. This parallel regulation of selective association with membranes and selective uptake by cells suggests a functional relationship. LDL, radiolabeled in its protein and in its lipid moiety, was incubated with liver plasma membranes. Besides LDL holo-particle receptor binding, also LDL cholesteryl esters were selectively associated with membranes. These data showed that selective association with membranes is not restricted to HDL but can occur from LDL as well. It is concluded that HDL3 as well as LDL cholesteryl esters can selectively be associated with hepatic plasma membranes, i.e., independent from particle binding. Results suggest an important role of the plasma membrane in the mechanism of selective cholesteryl ester uptake by the liver.

5'-Nucleotidase↗
PubMed1999

An alternating selection strategy for cloning phage display antibodies.

Phage display is a powerful technique that can be used to develop antibodies to target molecules. One approach for antibody phage display is to select phage from a large naive library of antibody immunoglobulin variable region fragments (Fv) expressed on the surface of phage. Phage that display antibody fragments of interest are selected by their ability to bind the target antigen immobilized on a solid support surface. A major difficulty often encountered with this approach is that phage that bind to additional antigens that are present during the phage selection steps are also selected. We have developed an alternating selection approach to minimize selection of unwanted phage. In the alternating selection approach, two selection methods are used. Each selection method contains different contaminating antigens. This approach was used to select phage that bind a phosphoryated form of the E47 transcription factor. Phage were selected based on their ability to bind a phospho-peptide in solution and alternatively a phospho-protein coated on a polyvinyl micortiter plate. This approach proved significantly better than selection with only one method. With one selection technique, 2 of 48 (4%) selected clones bound to the target antigen. With another selection technique, 15 of 48 (31%) selected clones bound to the target antigen. With alternating selection, 71 of 93 (76%) of the clones bound to the target antigen.

Antibodies↗
PubMed2006

The molecular signature of selection underlying human adaptations.

In the last decade, advances in human population genetics and comparative genomics have resulted in important contributions to our understanding of human genetic diversity and genetic adaptation. For the first time, we are able to reliably detect the signature of natural selection from patterns of DNA polymorphism. Identifying the effects of natural selection in this way provides a crucial piece of evidence needed to support hypotheses of human adaptation. This review provides a detailed description of the theory and analytical approaches used to detect signatures of natural selection in the human genome. We discuss these methods in relation to four classic human traits--skin color, the Duffy blood group, bitter-taste sensation, and lactase persistence. By highlighting these four traits we are able to discuss the ways in which analyses of DNA polymorphism can lead to inferences regarding past histories of selection. Specifically, we can infer the importance of specific regimes of selection (i.e. directional selection, balancing selection, and purifying selection) in the evolution of a trait because these different types of selection leave different patterns of DNA polymorphism. In addition, we demonstrate how these types of data can be used to estimate the time frame in which selection operated on a trait. As the field has advanced, a general issue that has come to the forefront is how specific demographic events in human history, such as population expansions, bottlenecks, and subdivision of populations, have also left a signature across the genome that can interfere with our detection of the footprint of selection at particular genes. Therefore, we discuss this general problem with respect to the four traits reviewed here, and describe the ways in which the signature of selection can be teased from a background signature of demographic history. Finally, we move from a discussion of analyses of selection motivated by a "candidate-gene" approach, in which a priori information led to the analysis of specific gene, to discussion of "genome-scanning" approaches that are directed at discovering new genes that have been under positive selection. Such scans can be designed to detect those genes that have been positively selected in our divergence from chimpanzees, as well as those genes that have been under selection as human populations have migrated, differentiated, and adapted to specific geographic environments. We predict that both approaches will be applied in the future, enabling a greater insight into human species-wide adaptations, as well as the specific adaptations of human populations.

Animals↗
PubMed2004

Selection and heritability of resistance to Bacillus thuringiensis subsp kurstaki and transgenic cotton in Helicoverpa armigera (Lepidoptera: Noctuidae).

Compared with an unselected susceptible population, a cotton bollworm, Helicoverpa armigera (Hübner), population selected for 22 generations with transgenic cotton leaves (modified Cry1A) in the laboratory developed 11.0-fold resistance to Cry1Ac (one single-protein product MVPII). Resistance to Bacillus thuringiensis Berliner subsp kurstaki (Btk) was selected for 22 generations with a 5.2-fold increase in LC50. The estimated realized heritabilities (h2) of resistance for transgenic-cotton- and Btk-selected populations were 0.1008 and 0.2341, respectively. This reflects the higher phenotypic variation in response to Cry1Ac in the transgenic-cotton-selected population. This variation may have been caused by differences in protein toxin levels expressed in different growth stages of the transgenic cotton. Because of the different slopes of the probit regression lines between Cry1Ac and Btk, the estimated realized h2 cannot be used visually to compare resistance development to Cry1Ac and Btk in H armigera. Thus, the response quotient (Q) of resistance was also estimated. The Q values of resistance for transgenic-cotton- and Btk-selected populations were 0.0763 and 0.0836, respectively. This showed that the rate of resistance development would be similar in both selection populations. This result indicates that the selection of resistance using transgenic cotton is different from that selected using the single toxin. Resistance risk to transgenic cotton and Btk in field populations was assessed assuming different pressures of selection by using the estimated h2. Assuming the h2 of resistance in a field population was half of the estimated h2, and the population received prolonged and uniform exposure to transgenic cotton or Btk causing >70% mortality in each generation, we predicted that resistance would increase 10-fold after <23 generations for Cry1Ac in transgenic cotton-selected-populations and after <21 generations for Btk in Btk-selected populations. Cross-resistance would be expected after <48 generations for Btk in transgenic-cotton-selected populations and after <21 generations for Cry1Ac in Btk-selected population. The results show that the potential to evolve resistance is similar in both transgenic-cotton- and Btk-selected populations, but that cross-resistance development to Btk is slower in transgenic-cotton-selected populations than cross-resistance development to Cry1Ac in Btk-selected populations.

Animals↗
PubMed1995

A genetical theory of species selection.

Species selection, differential rates of speciation or extinction resulting from species level characters, is often invoked as the main mechanism of macroevolution that is not simply an extension of microevolutionary processes. So long as we are careful in defining "species", the logic of species selection is sound. This does not mean, however, that this process can influence evolutionary dynamics under realistic conditions. The principal challenge to the efficacy of species selection as an evolutionary mechanism is the idea that selection between individuals within species will be so much more efficient as to swamp out any effects of selection between species. To assess this, a genetic model is constructed that includes simultaneous selection within and between species, and this is used to ask: under what conditions could species selection influence evolutionary dynamics, even in the face of opposing selection between individuals? The model shows that the efficacy of species selection is strongly determined by the time between speciation events (measured in individual generations), the mutation rate of the character under consideration, and the initial size of a newly formed reproductively isolated population. Data indicate that a few studied lineages have shown sufficiently high speciation rates to make species selection an important mechanism in the evolution of characters with mutation rates on the order of 10(-6) per generation. Quantitative characters, such as body size, generally change too readily for species selection to be relevant to their evolution. Complex characters, however, may be good candidates to be influenced by species selection. The interaction of selection within and between species can be subtle, with individual selection looking, from the standpoint of a species, very much like development of an individual. Furthermore, selection between individuals may be the main process assembling complex adaptations, while species selection allows them to persist over long periods of time.

Animals↗
PubMed1997

Genetic selection strategies: computer modeling.

There are four primary factors to consider in genetic selection strategies: 1) accuracy of selection, 2) selection intensity, 3) effective population size, and 4) mating system. Current theory indicates that optimum response to selection is achieved by maximizing the first three factors and using a mating systems that allows optimization of reproductive characteristics in dam lines and production characteristics in sire lines. However, with limited resources, compromises among the first three factors are needed. Simulations are useful for examining those compromises. Unrealistic simplifying assumptions are necessary for analytic theoretical results and thus do not address real world breeding problems. Using simulations, the relationship between selection accuracy, which is increased by use of family selection indices or Best Linear Unbiased Prediction (BLUP), and response to selection was examined. Results show that those procedures place a great restriction on effective population size, which offsets most of their advantage, i.e., there is too little emphasis on effective population size. A revision of the methodology and a reappraisal of the results of selection theory for optimization of genetic response is required. Another relationship that is of fundamental importance in breeding programs is that between selection intensity and effective population size. Analytical results for the additive case have been developed but have never been extended to heterotic traits. A gene level simulation program was developed to examine that relationship. Results show that the optimal selection strategy depends on the trait being selected. For additive traits and in the short term (20 generations), one should maximize selection intensity. For heterotic traits, an intermediate proportion (25% of each sex) gives optimal response. In all breeding strategies, primary attention must be given to the rate of inbreeding, which is increased by increasing either accuracy of selection or selection intensity. Inbreeding reduces response to selection in two ways. First, for both additive and nonadditive traits, inbreeding is a measure of the amount of random genetic drift that has occurred. Genetic drift causes loss of favorable alleles. Once lost, those alleles can never be recovered and thus genetic drift lowers the selection limit. Second, for heterotic traits, inbreeding results in a depression of the mean caused by directional dominance.

Animals↗
PubMed2002

Sexual selection fails to promote adaptation to a new environment.

Selection can be divided into sexual and nonsexual components. Some work finds that a component of sexual selection, adaptive female selection for good genes, can promote nonsexual fitness. Less studied is the benefit from sexual selection in toto, that is, when intra- and intersexual selection are both present and able to affect females directly and indirectly. Here an upper bound for the net benefit of sexual selection is estimated for Drosophila melanogaster. Replicate populations were allowed to adapt to low-grade thermal stress, with or with out the operation of sexual selection. Because proteins and lipids are highly sensitive to temperature, low-grade thermal stress will select broadly across the genome for alternative alleles. Such broad, directional selection for thermal tolerance should increase the measurable benefits of sexual selection far beyond that available under stabilizing selection. Sexual selection was removed by enforced monogamy without mate choice and retained by enforced polyandry (four males per female). After 36 generations of thermal stress exposure, there was substantial adaptation to the new environment (the net reproductive rate increased six standard deviations relative to thermal controls). However, sexual selection did not affect the rate of adaptation. Therefore, adaptive female selection for thermal tolerance either was insignificant or negated by other aspects of sexual selection, for example, male-induced female harm, which has been shown to diminish under monogamy. This experiment employed two parameters that reduced the opportunity for divergence in such harm: a truncated intersexual interaction period and strong directional selection for thermal tolerance. No divergence in male-induced harm was observed.

Adaptation, Physiological↗
PubMed2006

Selection on body size and sexual size dimorphism differs between host species in a seed-feeding beetle.

Sexual size dimorphism varies substantially among populations and species but we have little understanding of the sources of selection generating this variation. We used path analysis to study how oviposition host affects selection on body size in a seed-feeding beetle (Stator limbatus) in which males contribute large ejaculates (nuptial gifts) to females. Females use nutrients in these ejaculates for egg production. Male body size, which affects ejaculate size, affects female fecundity and is thus under fecundity selection similar in magnitude to the fecundity selection on female body size. We show that when eggs are laid on a host on which larval mortality is low (seeds of Acacia greggii) fecundity predicts fitness very well and fecundity selection is the major source of selection on both male and female adult size. In contrast, when eggs are laid on a host on which larval mortality is high (seeds of Parkinsonia florida) fecundity poorly predicts fitness such that fecundity selection is relaxed on both male and female size. However, because egg size affects larval mortality on this poor host (P. florida) there is selection on female size via the female size --> egg size --> fitness path; this selection via egg size offsets the reduction in fecundity selection on female, but not male, body size. Thus, differences in host suitability (due to differences in larval mortality) affect the relative importance of two sources of selection on adult body size; fecundity selection on both male and female body size is lower on the poor quality host (P. florida) relative to the high quality host (A. greggii) whereas selection on female body size via effects of egg size on offspring survival (body size --> egg size --> fitness) is greater on the poor quality host relative to the high quality host. Because selection via the egg size path affects only females the difference in larval survival between hosts shifts the relative magnitude of selection on female vs. male size. Researchers working on other study systems should be alerted to the possible importance of subtle, but consequential, indirect selection on their study organisms.

Animals↗
PubMed2006

The evolution of aging and age-related physical decline in mice selectively bred for high voluntary exercise.

We tested whether selective breeding for early-age high voluntary exercise behavior over 16 generations caused the evolution of lifelong exercise behavior, life expectancy, and age-specific mortality in house mice (Mus domesticus). Sixteenth-generation mice from four replicate selection lines and four replicate random-bred control lines were individually housed from weaning through death and divided between two activity treatments (either with or without running wheels). Thus, there were four treatment groups: selection versus control crossed with active versus sedentary. The effects of selective breeding on life expectancy and age-specific mortality differed between females and males. In females, sedentary selection mice had early and high initial adult mortality and thus the lowest increases in mortality with age. Active selection females had the lowest early adult mortality, had limited mortality during midlife, and exhibited rapid increases in mortality rates at the very end of life; thus, they had deferred senescence. Median life expectancy was greater for both groups of selection females than for the two complementary groups of control females. Like females, sedentary selection males had the highest early adult mortality, and slow but steadily increasing mortality over the entire lifetime. Unlike the active selection females, active control males had the lowest mortality across the lifespan (until the end of life). Interestingly, the males with the lowest median life expectancy were those in the active selection treatment group. In both sexes, running (km/week) decreased over the lifetime to very low and virtually equivalent levels at the end of life in control and selection mice. Overall, these results demonstrate an evolutionary cost of selective breeding for males, regardless of exercise level, but a benefit for females when they have an outlet for the up-selected behavior. We conclude that correlated evolution of senescence occurs in mice selectively bred for high voluntary wheel running; exercise per se is beneficial for control mice of both sexes, but the impact on the effect of selection depends on sex; and the behavioral effect of exercise selection at an early age declines throughout the life span, which demonstrates decreasing genetic correlations over age for the genes involved in increased exercise.

Aging↗
PubMed2005

The value of genetic information in selecting dairy replacements.

The objective of this study was to empirically determine the economic value of genetic information in the selection of dairy replacements, and assess whether this value was sufficient to prompt producers to select replacements on this basis. The data set consisted of 1982 Michigan Holstein replacements in 115 herds. Each herd had a minimum of 10 replacements that were born in the last 6 months of 1992 and calved within the last 6 months of 1994. The data for each replacement included the estimated breeding value (EBV) for milk at the beginning and end of the rearing period, and the estimated lifetime profit corrected for the opportunity cost of postponed replacement (ELPCOC). The replacement selection decision for a profit-maximizing dairy producer selecting 70 or 80% of the replacements was modeled. We modeled three methods of selection: genetic, random and ex poste. Genetic selection was evaluated using the EBV milk available at the beginning or end of the rearing period. For each herd, the profit associated with each of the three methods of selection was simulated. The value of the genetic information and perfect information were the differences in herd profits of genetic selection and ex poste selection relative to random selection, respectively. The difference in value of the genetic information between the end of the rearing period and the beginning of the rearing period was the increase in value of the genetic information due to updating. The value of information was calculated as the average herd profit per replacement. The value of the genetic information ranged from 22 dollars/replacement to 30 dollars/replacement and was statistically greater than zero at a 95% confidence level. It is unclear whether this value is sufficient to prompt producers to select replacements on the basis of EBV milk as has been recommended. The negative value of EBV milk (from the end of the rearing period when selecting 80% of the replacements) for 32 herds was consistent with the noisiness of the genetic estimates as messages of ELPCOC. The increased value of the genetic information due to updating was approximately 5 dollars/replacement. This increased value is likely insufficient to warrant delaying replacement selection decisions solely to obtain the updated information. The value of EBV milk was approximately 4 dollars/replacement higher when selecting 70% of the replacements versus 80%. The genetic information captured between 15% (selecting 70% at the beginning of the rearing period) and 20% (selecting 80% at the end of the rearing period) of the value of perfect information.

Animals↗
PubMed2001

Tumor cell depletion of peripheral blood progenitor cells using positive and positive/negative selection in metastatic breast cancer.

BACKGROUND: The clinical relevance of tumor cell purging of hematopoietic progenitor cell grafts has yet to be conclusively determined. Therefore, in addition to the demonstration that a method for graft purification is capable of removing an adequate number of tumor cells, it is critical that the procedure has as benign an impact upon the hematopoietic repopulating potential of the graft as possible. We evaluated tumor cell depletion, recovery of CD34(+) cells and post transplant engraftment kinetics as accepted measures of the effectiveness of an immunomagnetic bead (positive and positive/negative) purging methodology. METHODS: The patients received either positive selection (CD34 selection alone) or a combination of positive and negative (CD34 selection followed by breast cancer cell depletion) using the Isolex 300 (automated and semiautomated) devices. Immunocytochemistry was used to determine the degree of breast cancer cell contamination before and after the selection procedures to determine the efficacy of the procedure. CD34 enumeration was employed to evaluate the recovery and purity of the CD34-selected cellular products and engraftment indices (days to absolute neutrophil count (ANC) recovery and platelet count (Plt) recovery and transfusion requirements) were evaluated to determine the safety of the procedure. RESULTS: A total of 130 aphereses was performed on 101 patients. Ten pairs of collections were pooled before selection to increase the likelihood of achieving CD34 dose goals after selection. In all, 100 positive selections and 20 positive/negative selections were performed. Of the 10 (10.4%) ICC-positive preselection samples, 2 products showed persistent contamination after processing. The majority of patients (85.4%) required one selection procedure to achieve an adequate CD34(+) selected cell dose. Median CD34(+) cell recovery was > 50% for positive selection procedures and > 60% for the positive/negative procedures. The dose of CD34(+) cells infused ranged from 0.76 x 10(6) CD34(+) cells/kg to 27.7 x 106 CD34(+) cells/kg. There were no significant delays in neutrophil or platelet recovery or infections between any of the treatment groups. DISCUSSION: CD34 selection alone or in combination with negative selection can result in a significant reduction of contaminating tumor cells in the peripheral blood progenitor cell autograft. Although there was one engraftment failure with the CD34-positive selected cells, transplantation of the selected products after high-dose chemotherapy for metastatic breast cancer did not result in a clinically significant delay in the hematopoietic reconstitutive capacity of the autografts.

Adult↗
PubMed1988

Direction-selective adaptation in simple and complex cells in cat striate cortex.

1. The selectivity of adaptation to unidirectional motion was examined in neurons of the cat striate cortex. Following prolonged stimulation with a unidirectional high-contrast grating, the responsivity of cortical neurons was reduced. In many units this decrease was restricted to the direction of prior stimulation. This selective adaptation produced changes in the degree of direction selectivity of the cortical units (as measured by the ratio of the response to motion in the preferred direction to that in the nonpreferred direction). 2. The initial strength of the directional preference of a given cortical unit did not determine the degree of direction-selective adaptation. Indeed, even non-direction-selective units could exhibit pronounced direction-selective adaptation. The degree of direction-selective adaptation was also independent of the overall decrease in responsivity during adaptation. 3. There was no difference between simple and complex cells in the total amount of adaptation observed. The selectivity of the adaptation, however, did differ between these two cell types. As a group, simple cells showed significant direction-selective adaptation, whereas complex cells did not. The directional preference of most simple cells decreased following preferred direction adaptation and many highly direction selective simple cells became non-direction selective. In addition, simple cells became significantly more direction selective following nonpreferred direction adaptation. 4. Some complex cells also demonstrated direction-selective adaptation. There was, however, much more variability among complex cells than simple cells. Some complex cells actually increased direction selectivity following preferred direction adaptation. These differences between simple and complex cells suggest that changes in direction selectivity following unidirectional adaptation are not due to simple neuronal fatigue of the unit being recorded, but depend on selective adaptation of afferent inputs to the unit. 5. The spontaneous activity of many cortical neurons decreased following preferred direction adaptation but increased following adaptation in the nonpreferred direction. The response to a stationary grating also decreased following preferred direction adaptation. However, there was very little change in the response to a stationary grating following adaptation in the nonpreferred direction.

Adaptation, Physiological↗
PubMed1998

Strobe rearing reduces direction selectivity in area 17 by altering spatiotemporal receptive-field structure.

Strobe rearing reduces direction selectivity in area 17 by altering spatiotemporal receptive-field structure. J. Neurophysiol. 80: 2991-3004, 1998. Direction selectivity in simple cells of cat area 17 is linked to spatiotemporal (S-T) receptive-field structure. S-T inseparable receptive fields display gradients of response timing across the receptive field that confer a preferred direction of motion. Receptive fields that are not direction selective lack gradients; they are S-T separable, displaying uniform timing across the field. Here we further examine this link using a developmental paradigm that disrupts direction selectivity. Cats were reared from birth to 8 mo of age in 8-Hz stroboscopic illumination. Direction selectivity in simple cells was then measured using gratings drifting at different temporal frequencies (0.25-16 Hz). S-T structure was assessed using stationary bars presented at different receptive-field positions, with bar luminance being modulated sinusoidally at different temporal frequencies. For each cell, plots of response phase versus bar position were fit by lines to characterize S-T inseparability at each temporal frequency. Strobe rearing produced a profound loss of direction selectivity at all temporal frequencies; only 10% of cells were selective compared with 80% in normal cats. The few remaining directional cells were selective over a narrower than normal range of temporal frequencies and exhibited weaker than normal direction selectivity. Importantly, the directional loss was accompanied by a virtual elimination of S-T inseparability. Nearly all cells were S-T separable, like nondirectional cells in normal cats. The loss was clearest in layer 4. Normally, inseparability is greatest there, and it correlates well (r = 0.77) with direction selectivity; strobe rearing reduced inseparability and direction selectivity to very low values. The few remaining directional cells were inseparable. In layer 6 of normal cats, most direction-selective cells are only weakly inseparable, and there is no consistent relationship between the two measures. However, after strobe rearing, even the weak inseparability was eliminated along with direction selectivity. The correlated changes in S-T structure and direction selectivity were confirmed using conventional linear predictions of directional tuning based on responses to counterphasing bars and white noise stimuli. The developmental changes were permanent, being observed up to 12 yr after strobe rearing. The deficits were remarkably specific; strobe rearing did not affect spatial receptive-field structure, orientation selectivity, spatial or temporal frequency tuning, or general responsiveness to visual stimuli. These results provide further support for a critical role of S-T structure in determining direction selectivity in simple cells. Strobe rearing eliminates directional tuning by altering the timing of responses within the receptive field.

Animals↗
PubMed2005

Impact of genetic selection on management of boar replacement.

Boars in an artificial insemination centre have been selected for their superior genetic potential, with 'superior' being defined as having traits the customer wants transmitted to his herd. The ability to meet the customers' needs depends on the heritability of the trait, the geneticist's success in devising a selection scheme for the trait in balance with other economically important traits, and the boar's ability to produce sperm that can fertilise oocytes. Genetic evaluation research over the past 20 years has greatly increased the number of traits for which a boar can be selected: currently in the Canadian national program, these include age at 100 kg, backfat at 100 kg, feed efficiency, lean yield and litter size. In the near future, traits that are very likely to be added to this selection list include piglet survival, marbling, loin eye area and structure traits. In Canada, sires are ranked on two estimated breeding value (EBV) indices; one, focused on development of terminal sire lines, is based on the growth and yield traits and another, primarily focused on maternal line development, de-emphasises these traits and incorporates litter size. Boars that are in Canadian AI centres because of their excellent growth traits are typically in the top 5-10% of the national population for terminal sire line index, but they may be only average or substandard for litter size. Conversely, boars selected to be in the top 5-10% for conveying such reproductive traits as litter size may only be in the top 33% for growth traits. The more offspring from a superior boar in either of these indices, the faster the population average for the trait improves. The original sire gets knocked out of the elite group, is culled and replaced by a higher ranked young boar from the now improved general population. Although genetic superiority should govern an AI centre's selection and culling of boars, decision-making in real life is seldom that simple. Selection criteria may be contradictory as above, or a boar with truly superior traits may be excluded because a newly-developed molecular genetics test determines he carries an undesirable gene such as PSS, RN or others being developed. Selection for terminal sire or maternal line traits can ignore important practical factors that affect an AI centre--boars with superior genetics may not produce good semen because skeletal or penile problems prevent ejaculation, or because sperm production is poor due to a genetic flaw, disease, or some other cause. Interestingly, selection pressure for one trait may inadvertently select for a trait that is linked but whose linkage is unrecognised, and such unintentionally selected genes could benefit, harm, or have no effect on production traits. An AI centre serving a variety of customers must select boars in anticipation of their customers' needs (including new, foreign and niche markets). A centre should also review its genetic evaluation results and progeny records, both to critique its own selection success and to try to detect unexpected linkages. Finally, an AI centre needs to predict its own future, selecting not just for production traits for the swine producer, but also for factors that enhance the centre's efficiency including boar conformation and temperament, and sperm quantity, quality and hardiness. Can we select for efficiency? Our colleagues in dairy cattle AI evaluate bull performance--should the swine industry consider evaluation of male fertility traits?

Animals↗
PubMed2001

Strength and tempo of directional selection in the wild.

Directional selection is a major force driving adaptation and evolutionary change. However, the distribution, strength, and tempo of phenotypic selection acting on quantitative traits in natural populations remain unclear across different study systems. We reviewed the literature (1984-1997) that reported the strength of directional selection as indexed by standardized linear selection gradients (beta). We asked how strong are viability and sexual selection, and whether strength of selection is correlated with the time scale over which it was measured. Estimates of the magnitude of directional selection (absolute value of beta) were exponentially distributed, with few estimates greater than 0.50 and most estimates less than 0.15. Sexual selection (measured by mating success) appeared stronger than viability selection (measured by survival). Viability selection that was measured over short periods (days) was typically stronger than selection measured over longer periods (months and years), but the strength of sexual selection did not vary with duration of selection episodes; as a result, sexual selection was stronger than viability selection over longer time scales (months and years), but not over short time scales (days).

Animals↗
PubMed1972

Studies on the scutellar bristles of Drosophila melanogaster. II. Long-term selection for high bristle number in the Oregon RC strain and correlated responses in abdominal chaetae.

Results are presented of 135 generations of selection for high scutellar bristle number in two lines M and M3 derived from the same original mating of one female with 5 bristles by one male with 4 bristles, the latter being the wild-type canalised phenotype. Results are also given of two relaxed lines per line and of a reselection line M2 derived from the first relaxed line of line M which had regressed almost to base population level. The effect of introducing the sc(1) allele into the M and M3 selected backgrounds was studied at generations 39-44. At the end of selection the effect of an extra dose of sc(+) was also studied in males of all selected backgrounds. The correlated responses in abdominal bristles were followed in all lines.-Considering their common origin, the selection lines differed markedly in pattern of scutellar response and in most other aspects observed, namely correlated responses in abdominals and p.c. scutellars, sex differences, and behaviour on relaxation. Selection limits for scutellar bristles in lines M and M2 were equal to or greater than the most extreme reported in the literature.-The probit span of the canalised 4 bristle class decreased in each selection line as the mean scutellar bristle number increased, and increased again in the relaxed lines as the mean bristle number decreased. In the context of an hypothesis that canalisation at 4 bristle is due to regulation of the scute locus, this result is now interpreted as being due mainly to selection for poor regulators of sc(+), in contrast to a previous interpretation that only the minor gene background was altered by selection, the canalisation (regulation) genotype not being affected.-Introducing the sc(1) allele into the selected backgrounds M and M3 showed a reduced effect on sc(1) flies compared with sc(+) flies, and an interaction of sc(1) and sc(+) with selected background. sc(1) flies had about the same number of bristles in both backgrounds though the mean of sc(+) flies in line M was about 3sigma higher than in line M3. Dominance of sc(+) to sc(1) was reduced slightly in M3. However, the effect of an extra dose of sc(+) at the end of selection was about the same as in unselected in all lines, so the first or dominance level of regulation of the scute locus was not significantly affected by selection, though the second or canalisation level of regulation was.-A large positive correlated response in abdominal bristles occurred in all lines. The response in line M was about twice that in M2 and M3 and was in fact as large as can be obtained from direct selection on abdominals. In line M some genes may have been selected with a proportionately greater effect on abdominals than on scutellars. This is supported by the further observation in line M that the abdominal scores of flies with particular scutellar bristles scores increased as the scutellar mean increased. An attempt was made to apply to these results Rendel's (1962) model of competition between scutellars and abdominals for common bristle-making resources. This could not be done satisfactorily mainly because the assumptions in the model about the similarity of effects in scute and wild-type flies were not met in the present material.

Abdomen↗
PubMed2000

The evolution of genetic canalization under fluctuating selection.

If the direction of selection changes from generation to generation, the ability to respond to selection is maladaptive: the response to selection in one generation leads to reduced fitness in the next. Because the response is determined by the amount of genetic variance expressed at the phenotypic level, rapidly fluctuating selection should favor modifier genes that reduce the phenotypic effect of alleles segregating at structural loci underlying the trait. Such reduction in phenotypic expression of genetic variation has been named "genetic canalization." I support this argument with a series of two- and multilocus models with alternating linear selection and Gaussian selection with fluctuating optimum. A canalizing modifier gene affects the fitness of its carriers in three ways: (1) it reduces the phenotypic consequences of genetic response to previous selection; (2) it reduces the genetic response to selection, which is manifested as linkage disequilibrium between the modifier and structural loci; and (3) it reduces the phenotypic variance. The first two effects reduce fitness under directional selection sustained for several generations, but improve fitness when the direction of selection has just been reversed. The net effect tends to favor a canalizing modifier under rapidly fluctuating selection regimes (period of eight generations or less). The third effect improves fitness of the modifier allele if the fitness function is convex and reduces it if the function is concave. Under fluctuating Gaussian selection, the population is more likely to experience the concave portion of the fitness function when selection is stronger. Therefore, only weak to moderately strong fluctuating Gaussian selection favors genetic canalization. This paper considerably broadens the conditions that favor genetic canalization, which so far has only been postulated to evolve under long-term stabilizing selection.

Alleles↗