Evolutionary implications of host–pathogen specificity: the fitness consequences of host life history traits

Pathogens and parasites can be strong agents of selection, and often exhibit some degree of genetic specificity for individual host strains. Here we show that this host–pathogen specificity can affect the evolution of host life history traits. All else equal, evolution should select for genes that increase individuals' reproduction rates or lifespans (and thus total reproduction per individual). Using a simple host–pathogen model, we show that when the genetic specificity of pathogen infection is low, host strains with higher reproduction rates or longer lifespans drive slower-reproducing or shorter-lived host strains to extinction, as one would expect. However, when pathogens exhibit specificity for host strains with different life history traits, the evolutionary advantages of these traits can be greatly diminished by pathogen-mediated selection. Given sufficient host–pathogen specificity, pathogen-mediated selection can maintain polymorphism in host traits that are correlated with pathogen resistance traits, despite large intrinsic fitness differences among host strains. These results have two important implications. First, selection on host life history traits will be weaker than expected, whenever host fitness is significantly affected by genotype-specific pathogen attack. Second, where polymorphism in host traits is maintained by pathogen-mediated selection, preserving the genetic diversity of host species may require preserving their pathogens as well. This revised version was published online in November 2006 with corrections to the Cover Date.     

Genetic analysis of extended lifespan in Drosophila melanogaster III. On the relationship between artificially selected and wild stocks

Adult lifespans, age‐specific survival, age‐specific mortality, survival times on paraquat, and survival times on DDT were assayed in seven lines of Drosophila melanogaster, including two genetically heterogeneous wild lines recently collected from nature, and three inbred and recombinant inbred lines derived from an artificial selection experiment for increased lifespan. Survival on paraquat is positively correlated with adult lifespan. DDT resistance is uncorrelated with either paraquat resistance or lifespan. The wild lines are unexceptional with respect to average lifespan, paraquat resistance, age‐specific survivorship, and leveling off of mortality rates at advanced ages, but have high levels of DDT resistance. Cluster analysis groups the wild lines with three unselected laboratory stocks in one cluster, while two long‐lived elite recombinant inbred lines form a second cluster. Long‐lived laboratory‐adapted lines are quantitatively differentiated from the wild stocks, both with respect to average adult lifespans and resistance to an oxidizing agent. We reject the ‘recovery’ hypothesis, which proposes that Drosophila artificially selected for long life have phenotypes that merely recover the wild state. This revised version was published online in July 2006 with corrections to the Cover Date.     

Protein restriction and branched‐chain amino acid restriction promote geroprotective shifts in metabolism

The proportion of humans suffering from age‐related diseases is increasing around the world, and creative solutions are needed to promote healthy longevity. Recent work has clearly shown that a calorie is not just a calorie—and that low protein diets are associated with reduced mortality in humans and promote metabolic health and extended lifespan in rodents. Many of the benefits of protein restriction on metabolism and aging are the result of decreased consumption of the three branched‐chain amino acids (BCAAs), leucine, isoleucine, and valine. Here, we discuss the emerging evidence that BCAAs are critical modulators of healthy metabolism and longevity in rodents and humans, as well as the physiological and molecular mechanisms that may drive the benefits of BCAA restriction. Our results illustrate that protein quality—the specific composition of dietary protein—may be a previously unappreciated driver of metabolic dysfunction and that reducing dietary BCAAs may be a promising new approach to delay and prevent diseases of aging.     

江西九连山常绿阔叶林主要树种叶建成消耗的比较

叶建成消耗（或叶构建消耗）是指构建单位质量（面积）叶片所需要的葡萄糖当量,是植物碳收获过程中必要的成本投资．反映植物在叶片水平上的能量和碳分配策略。基于热值、灰份含量、叶氮含量和比叶面积的测定,估算了江西九连山常绿阔叶林16个主要树种的叶建成消耗,分析比较了不同树种、不同叶寿命、比叶面积和冠层不同部位对叶建成消耗的影响。结果表明,热值组分的变化对单位质量叶建成消耗的影响最大;树种间单位质量叶建成消耗的差异不大（变化率仅3％～6％）,但其单位面积叶建成消耗则存在显著差异（变化率达27％～28％）;热值和单位质量叶建成消耗与比叶面积密切相关,而比叶面积随取样高度呈显著的降低趋势,最终导致热值及单位质量和单位面积的叶建成消耗均随取样高度呈显著的增加趋势。研究结果指出：（1）较高的叶建成消耗是由于植物叶片内含有较高能量投资的化学组分;（2）叶建成消耗和比叶面积随取样高度的变化综合反映了植物对光照和水分资源垂直梯度变化的可塑性适应。     

Small trees,big problems: Comparative leaf function under extreme edaphic stress

Premise of the Study

The pygmy forest, a plant community of severely stunted conifers and ericaceous angiosperms, occurs on patches of highly acidic, nutrient‐poor soils along the coast of Northern California, USA. This system is an excellent opportunity to study the effect of severe nutrient deficiency on leaf physiology in a naturally‐occurring ecosystem. In this study, we seek to understand the physiological mechanisms stunting the plants' growth and their implications for whole plant function.

Methods

We measured 14 traits pertaining to leaf photosynthetic function or physical structure on seven species. Samples were taken from the pygmy forest community and from conspecifics growing on higher‐nutrient soils, where trees may grow over 30 m tall.

Key Results

Pygmy plants of most species maintained similar area‐based photosynthetic and stomatal conductance rates to conspecific controls, but had lower specific leaf area (leaf area divided by dry weight), lower percent nitrogen, and less leaf area relative to xylem growth. Sequoia sempervirens, a species rare in the pygmy forest, had a categorically different response from the more common plants and had remarkably low photosynthetic rates.

Conclusions

Pygmy plants were not stunted by low photosynthetic rates on a leaf‐area basis; instead, several species had restricted whole‐plant photosynthesis due to low leaf area production. Pygmy plants of all species showed signs of greater carbon investment in their leaves and higher production of nonphotosynthetic leaf tissue, further contributing to slow growth rates.     

Strong dominance of functional alleles over gene deletions in both intensely growing and deeply starved yeast cells

Previous studies with diploid yeast have shown that the deletion of one allele at a single locus typically has little impact on fitness under conditions promoting fast growth. Here, we confirm and quantify this finding. The strong dominance of functional over nonfunctional alleles is predicted by the metabolic control theory which assumes that the cell is a system of metabolic fluxes and that the total metabolic rate is equivalent to fitness. To test whether these requirements are critical, we tested dominance under conditions of long‐term starvation when metabolism is low and thus the metabolic activities of proteins are likely inadequate or imbalanced. More fundamentally, the central assumption of the model, that high metabolic rate translates into high fitness, appears implausible. Contrary to these conjectures, we found that the mean rate of survival of starving heterozygotes was affected only slightly more than was the mean rate of growth under good conditions. Under none of the two treatments the central prediction of the model, that fitness of heterozygous strains is higher for the enzymatic proteins than for nonenzymatic ones, was confirmed. Our data add to growing uncertainty whether the metabolic control theory is sufficient to explain the remarkable ubiquity of strong genetic dominance.     

Leaf lifespan as a determinant of leaf structure and function among 23 amazonian tree species

Summary The relationships between resource availability, plant succession, and species' life history traits are often considered key to understanding variation among species and communities. Leaf lifespan is one trait important in this regard. We observed that leaf lifespan varies 30-fold among 23 species from natural and disturbed communities within a 1-km radius in the northern Amazon basin, near San Carlos de Rio Negro, Venezuela. Moreover, leaf lifespan was highly correlated with a number of important leaf structural and functional characterisues. Stomatal conductance to water vapor (g) and both mass and area-based net photosynthesis decreased with increasing leaf lifespan (r²=0.74, 0.91 and 0.75, respectively). Specific leaf area (SLA) also decreased with increasing leaf lifespan (r²=0.78), while leaf toughness increased (r²=0.62). Correlations between leaf lifespan and leaf nitrogen and phosphorus concentrations were moderate on a weight basis and not significant on an area basis. On an absolute basis, changes in SLA, net photosynthesis and leaf chemistry were large as leaf lifespan varied from 1.5 to 12 months, but such changes were small as leaf lifespan increased from 1 to 5 years. Mass-based net photosynthesis (A/mass) was highly correlated with SLA (r²=0.90) and mass-based leaf nitrogen (N/mass) (r²=0.85), but area-based net photosynthesis (A/area) was not well correlated with any index of leaf structure or chemistry including N/area. Overall, these results indicate that species allocate resources towards a high photosynthetic assimilation rate for a brief time, or provide resistant physical structure that results in a lower rate of carbon assimilation over a longer time, but not both.     

DNA replication stress-induced loss of reproductive capacity in S. cerevisiae and its inhibition by caloric restriction

In many organisms, attenuation of growth signaling by caloric restriction or mutational inactivation of growth signaling pathways extends lifespan and protects against cancer and other age-related diseases. The focus of many efforts to understand these effects has been on the induction of oxidative stress defenses that inhibit cellular senescence and cell death. Here we show that in the model organism S. cerevisiae, growth signaling induces entry of cells in stationary phase into S phase in parallel with loss of reproductive capacity, which is enhanced by elevated concentrations of glucose. Overexpression of RNR1 encoding a ribonucleotide reductase subunit required for the synthesis of deoxynucleotide triphosphates and DNA replication suppresses the accelerated loss of reproductive capacity of cells cultured in high glucose. The reduced reproductive capacity of these cells is also suppressed by excess threonine, which buffers dNTP pools when ribonucleotide reductase activity is limiting. Caloric restriction or inactivation of the AKT homolog Sch9p inhibits senescence and death in stationary phase cells caused by the DNA replication inhibitor hydroxyurea or by inactivation of the DNA replication and repair proteins Sgs1p or Rad27p. Inhibition of DNA replication stress represents a novel mechanism by which caloric restriction promotes longevity in S. cerevisiae. A similar mechanism may promote longevity and inhibit cancer and other age-related diseases in humans.