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

I. the origin of species by means of natural selection

Reactive O2 species (ROS) are produced in both unstressed and stressed cells. Plants have welldeveloped defence systems against ROS, involving both limiting the formation of ROS as well as instituting its removal. Under unstressed conditions, the formation and removal of O2 are in balance. However, the defence system, when presented with increased ROS formation under stress conditions, can be overwhelmed. Within a cell, the superoxide dismutases (SODs) constitute the first line of defence against ROS. Specialization of function among the SODs may be due to a combination of the influence of subcellular location of the enzyme and upstream sequences in the genomic sequence. The commonality of elements in the upstream sequences of Fe, Mn and CuuZn SODs suggests a relatively recent origin for those regulatory regions. The differences in the upstream regions of the three FeSOD genes suggest differing regulatory control which is borne out in the research literature. The finding that the upstream sequences of Mn and peroxisomal CuuZn SODs have three common elements suggests a common regulatory pathway. The tools are available to dissect further the molecular basis for antioxidant defence responses in plant cells. SODs are clearly among the most important of those defences, when coupled with the necessary downstream events for full detoxification of ROS.

Role of superoxide dismutases (SODs) in controlling oxidative stress in plants

Species range limits involve many aspects of evolution and ecology, from species distribution and abundance to the evolution of niches. Theory suggests myriad processes by which range limits arise, including competitive exclusion, Allee effects, and gene swamping; however, most models remain empirically untested. Range limits are correlated with a number of abiotic and biotic factors, but further experimentation is needed to understand underlying mechanisms. Range edges are characterized by increased genetic isolation, genetic differentiation, and variability in individual and population performance, but evidence for decreased abundance and fitness is lacking. Evolution of range limits is understudied in natural systems; in particular, the role of gene flow in shaping range limits is unknown. Biological invasions and rapid distribution shifts caused by climate change represent large-scale experiments on the underlying dynamics of range limits. A better fusion of experimentation and theory will advance our understanding of the causes of range limits.

https://faculty.ucmerced.edu/sites/default/files/jasonsexton.com/files/sexton_2009_annualreviewofecologyevolutionandsystematics.pdf

Evolution and Ecology of Species Range Limits

Understanding how landscape characteristics affect biodiversity patterns and ecological processes at local and landscape scales is critical for mitigating effects of global environmental change. In this review, we use knowledge gained from human-modified landscapes to suggest eight hypotheses, which we hope will encourage more systematic research on

/pdf/landscape-moderation-of-biodiversity-patterns-and-processes-pokn3pbbbx.pdf

Landscape moderation of biodiversity patterns and processes - eight hypotheses

There is growing interest in quantifying genetic population structure across the geographical ranges of species to understand why species might exhibit stable range limits and to assess the conservation value of peripheral populations. However, many assertions regarding peripheral populations rest on the long-standing but poorly tested supposition that peripheral populations exhibit low genetic diversity and greater genetic differentiation as a consequence of smaller effective population size and greater geographical isolation relative to geographically central populations. We reviewed 134 studies representing 115 species that tested for declines in within-population genetic diversity and/or increases in among-population differentiation towards range margins using nuclear molecular genetic markers. On average, 64.2% of studies detected the expected decline in diversity, 70.2% of those that tested for it showed increased differentiation and there was a positive association between these trends. In most cases, however, the difference in genetic diversity between central and peripheral population was not large. Although these results were consistent across plants and animals, strong taxonomic and biogeographical biases in the available studies call for a cautious generalization of these results. Despite the large number of studies testing these simple predictions, very few attempted to test possible mechanisms causing reduced peripheral diversity or increased differentiation. Almost no study incorporated a phylogeographical framework to evaluate historical influences on contemporary genetic patterns. Finally, there has been little effort to test whether these geographical trends in putatively neutral variation at marker loci are reflected by quantitative genetic trait variation, which is likely to influence the adaptive potential of populations across the geographical range.

Genetic variation across species' geographical ranges: the central-marginal hypothesis and beyond

Many species’ range limits (RL) occur across continuous environmental gradients without obvious barriers imposing them. Such RL are expected to reflect niche limits (NL) and thus to occur where populations cease to be self-sustaining. Transplant experiments comparing fitness within and beyond species’ ranges can test this hypothesis, but interpretive power depends strongly on experimental design. We first identify often overlooked aspects of transplant design that are critical to establishing the causes of RL, especially incorporating transplant sites at, and source populations from, the range edge. We then conduct a meta-analysis of published beyond-range transplant experiments ( tests). Most tests (75%) found that performance declined beyond the range, with the strongest declines detected when the measure of performance was lifetime fitness (83%), suggesting that RL commonly involve niche constraints (declining habitat quality). However, only 46% supported range limits occurring at NL; 26% (most...

https://www.jstor.org/stable/10.1086/674525

Are Species’ Range Limits Simply Niche Limits Writ Large? A Review of Transplant Experiments beyond the Range

It is widely accepted that species are most abundant at the center of their geographic ranges and become progressively rarer toward range limits. Although the abundant center model (ACM) has rarely been tested with range-wide surveys, it influences much thinking about the ecology and evolution of species' distributions. We tested ACM predictions using two unrelated but ecologically similar plants, Camissonia cheiranthifolia and Abronia umbellata. We intensively sampled both throughout their one-dimensional distributions within the Pacific coastal dunes of North America, from northern Baja California, Mexico, to southern Oregon, USA. Data from >1100 herbarium specimens indicated that these limits have been stable for at least the last 100 years. Range-wide field surveys detected C. cheiranthifolia at 87% of 124 sites and A. umbellata at 54% of 113 sites, but site occupancy did not decline significantly toward range limits for either species. Permutation analysis did not detect a significant fit of geographical variation in local density to the ACM. Mean density did not correlate negatively with mean individual performance (plant size or number of seeds/plant), probably because both species occur at low densities. Although size and seeds per plant varied widely, central populations tended to have the highest values for size only. For C. cheiranthifolia, we observed asymmetry in the pattern of variation between the northern and southern halves of the range consistent with the long-standing prediction that range limits are imposed by different ecological factors in different parts of the geographical distribution. However, these asymmetries were difficult to interpret and likely reflect evolutionary differentiation as well as plastic responses to ecological variation. Both density and seeds per plant contributed to variation in seed production per unit area. In C. cheiranthifolia only, sites with highest seed production tended to occur at the range center, as predicted by the ACM and assumed by theory proposing that range limits evolve via antagonism between natural selection and gene flow.

Testing the abundant center model using range-wide demographic surveys of two coastal dune plants.

Dispersal may be favoured at geographic range edges by unstable population and metapopulation dynamics. However, dispersal may also evolve in response to geographic variation in other life-history traits, especially the mating system. Here, increased dispersal at range margins was tested for with a range-wide analysis of seed dispersal and mating system traits in Abronia umbellata, a plant endemic to Pacific coastal dunes of North America. Seeds disperse within winged anthocarps. Anthocarps from 34 populations varied widely in wing size (mass-corrected wing index). Wing index correlated negatively with threshold wind velocity for dispersal in wind tunnel tests, suggesting that wings facilitate tumbling over open sandy substrate. As predicted, wing index increased and threshold velocity decreased towards both range limits. Flower size, herkogamy and self-incompatibility declined towards range limits, indicating a shift to self-fertilization, and flower size and wing index correlated negatively. However, the increase in wing index towards range limits remained after statistically controlling flower size. These results are consistent with selection favouring dispersal at range margins. The evolutionary lability of dispersal across the range may affect the interaction between selection and gene flow in the establishment and maintenance of geographic range limits.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.2007.02349.x

Increased seed dispersal potential towards geographic range limits in a Pacific coast dune plant.

Expression of individual superoxide dismutase (SOD) transgenes improves environmental stress tolerance and biomass production in alfalfa (Medicago sativa L.). The objective of this study was to test the hypothesis that synergy exists between transgenic SOD stress-tolerance mechanisms, specifically that the simultaneous expression of two SOD transgenes confers greater benefit than the expression of a single SOD transgene. The hypothesis was tested through an evaluation of an F 1 family generated through a sexual cross of a hemizygous Mit-MnSOD plant and a hemizygous Chl-MnSOD-transgenic alfalfa plant which had previously been screened in field trials for improved persistence. Southern analyses revealed that the parents each had single insertion regions of the MnSOD cDNA and the inheritance followed the expected Mendelian ratios. Native PAGE gels and enzyme inhibition assays revealed the activity of the transgenic MnSOD isozymes. F 1 progeny containing either the Mit-MnSOD or the Chl-MnSOD transgene had significantly higher storage organ (crown + root) biomass compared to non-transgenic siblings. The joint expression of the transgenes resulted in a numerical increase in total SOD activity. However, F 1 progeny containing both transgenes had lower shoot and storage organ biomass compared to siblings having only one or the other transgene, a result that did not support the authors' hypothesis. It was postulated that a promoter with lower expression than 35S may be necessary if closely related transgenes are to be pyramided in the same plant.

/pdf/pyramiding-mn-superoxide-dismutase-transgenes-to-improve-1hk2p27w20.pdf

Karen E. Samis

Papers

Genetic variation across species' geographical ranges: the central-marginal hypothesis and beyond

Are Species’ Range Limits Simply Niche Limits Writ Large? A Review of Transplant Experiments beyond the Range

Testing the abundant center model using range-wide demographic surveys of two coastal dune plants.

Increased seed dispersal potential towards geographic range limits in a Pacific coast dune plant.

Pyramiding Mn‐superoxide dismutase transgenes to improve persistence and biomass production in alfalfa