So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

Ecological changes in the phenology and distribution of plants and animals are occurring in all well-studied marine, freshwater, and terrestrial groups These observed changes are heavily biased in the directions predicted from global warming and have been linked to local or regional climate change through correlations between climate and biological variation, field and laboratory experiments, and physiological research Range-restricted species, particularly polar and mountaintop species, show severe range contractions and have been the first groups in which entire species have gone extinct due to recent climate change Tropical coral reefs and amphibians have been most negatively affected Predator-prey and plant-insect interactions have been disrupted when interacting species have responded differently to warming Evolutionary adaptations to warmer conditions have occurred in the interiors of species’ ranges, and resource use and dispersal have evolved rapidly at expanding range margins Observed genetic shifts modulate local effects of climate change, but there is little evidence that they will mitigate negative effects at the species level

https://www.fws.gov/southwest/es/documents/R2ES/LitCited/LPC_2012/Parmesan_2006.pdf

Ecological and Evolutionary Responses to Recent Climate Change

Physiological and ecological constraints play key roles in the evolution of plant growth patterns, especially in relation to defenses against herbivores. Phenotypic and life history theories are unified within the growth-differentiation balance (GDB) framework, forming an integrated system of theories explaining and predicting patterns of plant defense and competitive interactions in ecological and evolutionary time. Plant activity at the cellular level can be classified as growth (cell division and enlargement) of differentiation (chemical and morphological changes leading to cell maturation and specialization). The GDB hypothesis of plant defense is premised upon a physiological trade-off between growth and differentiation processes. The trade-off between growth and defense exists because secondary metabolism and structural reinforcement are physiologically constrained in dividing and enlarging cells, and because they divert resources from the production of new leaf area. Hence the dilemma of plants: Th...

https://www.jstor.org/stable/pdfplus/2830650.pdf

The dilemma of plants: To grow or defend.

This article describes ways in which excess nitrogen from fossil fuel combustion may stress the biosphere. Nitrogen emissions can have a direct effect on air quality through both the oxidizing potential of nitrogen oxides and the role these compounds play in the formation of ozone. The complexity of these effects on water quality and on forest nutrition is discussed.

https://harvardforest.fas.harvard.edu/sites/harvardforest.fas.harvard.edu/files/publications/pdfs/Aber_BioScience_1989.pdf

Nitrogen saturation in northern forest ecosystems

Host plant quality is a key determinant of the fecundity of herbivorous insects. Components of host plant quality (such as carbon, nitrogen, and defensive metabolites) directly affect potential and achieved herbivore fecundity. The responses of insect herbivores to changes in host plant quality vary within and between feeding guilds. Host plant quality also affects insect reproductive strategies: Egg size and quality, the allocation of resources to eggs, and the choice of oviposition sites may all be influenced by plant quality, as may egg or embryo resorption on poor-quality hosts. Many insect herbivores change the quality of their host plants, affecting both inter- and intraspecific interactions. Higher-trophic level interactions, such as the performance of predators and parasitoids, may also be affected by host plant quality. We conclude that host plant quality affects the fecundity of herbivorous insects at both the individual and the population scale.

Host plant quality and fecundity in herbivorous insects

Our aim in this study was to determine how well phenotypic variation in foliar concentrations of carbon-based secondary compounds (CBSCs) in woody plants can be predicted on the basis of two resource-based hypotheses, i.e. the carbon-nutrient balance (CNB) and growth-differentiation balance (GDB) hypotheses. We conducted a meta-analysis of literature data with respect to responses of CBSCs, carbohydrates and nitrogen to six types of environmental manipulations (fertilization with nitrogen or phosphorus, shading, CO 2 enrichment, drought stress, ozone exposure). Plant responses to nitrogen fertilization, shading and CO 2 enrichment in terms of pooled CBSCs and carbohydrates were consistent with predictions made with the two hypotheses. However, among biosynthetically distinct groups of CBSCs only concentrations of phenylpropanoid-derived compounds changed as predicted; hydrolyzable tannins and terpenoids, in particular, were less responsive. Phosphorus fertilization did not affect concentrations of CBSC or primary metabolites. Plant responses to drought and ozone exposure presumably were driven by plant demands for particular types of compounds (osmolites in the case of drought and antioxidants in the case of ozone exposure) rather than by changes in resource availability. Based on the relative importance of the treatment effects, we propose a hierarchical model of carbon allocation to CBSCs. The model implies that CBSC production is determined by both resource availability and specific demand-side responses. However, these two mechanisms work at different hierarchical levels. The domain of the CNB and GDB hypotheses is at the high hierarchical levels, predicting the total amount of carbon that can be allocated to CBSCs. Predicting altered concentrations of individual CBSCs, i.e. low hierarchy levels, probably demands biosynthetically detailed models which also take into account the history of plant interactions with biotic and abiotic factors.

Regulation of Woody Plant Secondary Metabolism by Resource Availability: Hypothesis Testing by Means of Meta-Analysis

Global warming is predicted to cause distributional changes in organisms whose geographic ranges are controlled by temperature. We report a recent latitudinal and altitudinal expansion of the pine processionary moth, Thaumetopoea pityocampa, whose larvae build silk nests and feed on pine foliage in the winter. In north-central France (Paris Basin), its range boundary has shifted by 87 km northwards between 1972 and 2004; in northern Italy (Alps), an altitudinal shift of 110–230 m upwards occurred between 1975 and 2004. By experimentally linking winter temperature, feeding activity, and survival of T. pityocampa larvae, we attribute the expansions to increased winter survival due to a warming trend over the past three decades. In the laboratory we determined the minimum nest and night air temperatures required for larval feeding and developed a mechanistic model based on these temperature thresholds. We tested the model in a translocation experiment that employed natural temperature gradients as spatial an...

Expansion of geographic range in the pine processionary moth caused by increased winter temperatures

In this review, we test the hypothesis that abiotic stress increases the suitability of plants as food for herbivores. We conducted a meta-analysis that included 70 experimental studies in which insect performance was measured on woody plants subjected to water stress, pollution, and/or shading. Overall, plant stress had no significant effect on insect growth rate, fecundity, survival, or colonization density. We found great variation, however, in the magnitude and direction of insect responses among studies, most of which was related to insect feeding guild. In general, boring and sucking insects performed better on stressed plants, whereas plant stress adversely affected gall-makers and chewing insects. Reduction in performance of chewers was greater on stressed slow-growing plants than on stressed fast growers. Reproductive potential of sucking insects was increased by pollution but reduced by water stress. In some cases where sample sizes were small or the treatment periods short, apparent differences in insect responses to stress were probably artifacts due to inappropriate experimental design.

Insect performance on experimentally stressed woody plants: a meta-analysis.

Stressful Times for the Plant Stress: Insect Performance Hypothesis

Phratora vitellinae and Galerucella lineola are two leaf beetles that feed on willows (Salix) in central Sweden. When disturbed, larvae of P. vitellinae exude droplets of a defensive secretion from dorsal glands. The secretion contains salicyl aldehyde, the precursors for which are plant—derived salicylates like salicin and other chemically similar phenolic glycosides. In contrast, larvae of G. lineola lack dorsal glands, cannot convert salicylates to salicyl aldehyde, and no not produce such a defensive secretion. We evaluate the adult oviposition pattern, larval performances, and suceptibility to predators of these two beetles on three species of willows, two rich in salicylates (Salix fragilis and S. dasyclados) and one poor in salicylates (S. viminalis). Females of G. lineola oviposited preferentially on S. viminalis and S. fragilis where larvae developed rapidly and survival was high, and avoided S. dasyclados where larval performance was poor. Variation in leaf pubescence, toughness, and nitrogen content across willow species not explain observed patterns of larval performance. However, larval performance was inversely related to the concentration of salicylates in the three willows. By contrast, the oviposition preferences of P. vitellinae did not correspond well to patterns of larval performance. Salix fragilis was by far preferred choice of oviposition, and females rarely placed eggs on either S. viminalis or S. dasyclados. The performance of P. vitellinae larvae differed markedly on the two salicylate—rich willows; larvae survived best and developed most rapidly on S. fragilis, but mortality was high and development slow on S. dasyclados. Poor larval performance on S. dasyclados was associated with high concentrations of simple phenolic compounds other than salicylates. Despite differences in performance, larvae fed these two salicylate—rich willows produced a defensive secretion which effectively repelled coccinellid predators. Larvae fed the salicylate—poor S. viminalis survived and developed well in the absence of enemies, but lacked the salicylate—based defensive secretion and were readily eaten by coccinellids. Females of P. vitellinae apparently avoided S. viminalis as an oviposition host, not because larvae performed poorly, but because larvae were defenseless against some predators when raised on this salicylate—poor willow. We argued that herbivores such as P. vitellinae obtain enemy—free space on hosts from which they sequesters plant—derived allelochemicals for defense. The result appears to be dietary specialization on host—plant species that provide these precursors for defense. Other related herbivores like G. lineola that do not employ plant—derived defensive secretions, are not so constrained and feed on a wider range of host willows.

Stig Larsson

Papers

Regulation of Woody Plant Secondary Metabolism by Resource Availability: Hypothesis Testing by Means of Meta-Analysis

Expansion of geographic range in the pine processionary moth caused by increased winter temperatures

Insect performance on experimentally stressed woody plants: a meta-analysis.

Stressful Times for the Plant Stress: Insect Performance Hypothesis

Role of Enemy-Free Space and Plant Quality in Host-Plant Selection by Willow Beetles