Adaptive constraints at the range edge of a widespread and expanding invasive plant.

Identifying the factors that facilitate and limit invasive species' range expansion has both practical and theoretical importance, especially at the range edges. Here, we used reciprocal common garden experiments spanning the North/South and East/West range that include the North American core, intermediate and range edges of the globally invasive plant, Johnsongrass (Sorghum halepense) to investigate the interplay of climate, biotic interactions (i.e. competition) and patterns of adaptation. Our results suggest that the rapid range expansion of Johnsongrass into diverse environments across wide geographies occurred largely without local adaptation, but that further range expansion may be restricted by a fitness trade-off that limits population growth at the range edge. Interestingly, plant competition strongly dampened Johnsongrass growth but did not change the rank order performance of populations within a garden, though this varied among gardens (climates). Our findings highlight the importance of including the range edge when studying the range dynamics of invasive species, especially as we try to understand how invasive species will respond to accelerating global changes.

Suppression force-fields and diffuse competition: competition de-escalation is an evolutionarily stable strategy.

Competition theory is founded on the premise that individuals benefit from harming their competitors, which helps them secure resources and prevent inhibition by neighbours. When multiple individuals compete, however, competition has complex indirect effects that reverberate through competitive neighbourhoods. The consequences of such 'diffuse' competition are poorly understood. For example, competitive effects may dilute as they propagate through a neighbourhood, weakening benefits of neighbour suppression. Another possibility is that competitive effects may rebound on strong competitors, as their inhibitory effects on their neighbours benefit other competitors in the community. Diffuse competition is unintuitive in part because we lack a clear conceptual framework for understanding how individual interactions manifest in communities of multiple competitors. Here, I use mathematical and agent-based models to illustrate that diffuse interactions-as opposed to direct pairwise interactions-are probably the dominant mode of interaction among multiple competitors. Consequently, competitive effects may regularly rebound, incurring fitness costs under certain conditions, especially when kin-kin interactions are common. These models provide a powerful framework for investigating competitive ability and its evolution and produce clear predictions in ecologically realistic scenarios.

Effects of species interactions on the potential for evolution at species' range limits.

Species' ranges are limited by both ecological and evolutionary constraints. While there is a growing appreciation that ecological constraints include interactions among species, like competition, we know relatively little about how interactions contribute to evolutionary constraints at species' niche and range limits. Building on concepts from community ecology and evolutionary biology, we review how biotic interactions can influence adaptation at range limits by impeding the demographic conditions that facilitate evolution (which we term a 'demographic pathway to adaptation'), and/or by imposing evolutionary trade-offs with the abiotic environment (a 'trade-offs pathway'). While theory for the former is well-developed, theory for the trade-offs pathway is not, and empirical evidence is scarce for both. Therefore, we develop a model to illustrate how fitness trade-offs along biotic and abiotic gradients could affect the potential for range expansion and niche evolution following ecological release. The model shows that which genotypes are favoured at species' range edges can depend strongly on the biotic context and the nature of fitness trade-offs. Experiments that characterize trade-offs and properly account for biotic context are needed to predict which species will expand their niche or range in response to environmental change. This article is part of the theme issue 'Species' ranges in the face of changing environments (Part II)'.

Early Sibling Conflict May Ultimately Benefit the Family.

Relatives often interact differently with each other than with nonrelatives, and whether kin cooperate or compete has important consequences for the evolution of mating systems, seed size, dispersal, and competition. Previous research found that the larger of the size dimorphic seeds produced by the annual plant Aegilops triuncialis suppressed germination of their smaller sibs by 25%-60%. Here, we found evidence for kin recognition and sibling rivalry later in life among Aegilops seedlings that places seed-seed interactions in a broader context. In experiments with size dimorphic seeds, seedlings reduced the growth of sibling seedlings by ∼40% but that of nonsibling seedlings by ∼25%. These sequential antagonistic interactions between seeds and then seedlings provide insight into conflict and cooperation among kin. Kin-based conflict among seeds may maintain dormancy for some seeds until the coast is clear of more competitive siblings. If so, biotically induced seed dormancy may be a unique form of cooperation, which increases the inclusive fitness of maternal plants and offspring by minimizing competition among kin.

Rapid and repeated local adaptation to climate in an invasive plant.

Biological invasions provide opportunities to study evolutionary processes occurring over contemporary timescales. To explore the speed and repeatability of adaptation, we examined the divergence of life-history traits to climate, using latitude as a proxy, in the native North American and introduced European and Australian ranges of the annual plant Ambrosia artemisiifolia. We explored niche changes following introductions using climate niche dynamic models. In a common garden, we examined trait divergence by growing seeds collected across three ranges with highly distinct demographic histories. Heterozygosity-fitness associations were used to explore the effect of invasion history on potential success. We accounted for nonadaptive population differentiation using 11 598 single nucleotide polymorphisms. We revealed a centroid shift to warmer, wetter climates in the introduced ranges. We identified repeated latitudinal divergence in life-history traits, with European and Australian populations positioned at either end of the native clines. Our data indicate rapid and repeated adaptation to local climates despite the recent introductions and a bottleneck limiting genetic variation in Australia. Centroid shifts in the introduced ranges suggest adaptation to more productive environments, potentially contributing to trait divergence between the ranges.

Climatic niche shifts are common in introduced plants.

Our understanding of how climate influences species distributions and our ability to assess the risk of introduced species depend on the assumption that species' climatic niches remain stable across space and time. While niche shifts have been detected in individual invasive species, one assessment of ~50 plants in Europe and North America concluded that niche shifts were rare, while another concluded the opposite. These contradictory findings, limited in species number and geographic scope, leave open a need to understand how often introduced species experience niche shifts and whether niche shifts can be predicted. We found evidence of climatic niche shifts in 65-100% of 815 terrestrial plant species introduced across five continents, depending on how niche shifts were measured. Individual species responses were idiosyncratic, but we generally saw that niche shifts reflected changes in climate availability at the continent scale and were largest in long-lived and cultivated species. Smaller intercontinental niche shifts occurred within species' native ranges. Overall, the climatic niches of terrestrial plant species were not conserved as they crossed continents. These results have major consequences for applying environmental niche models to assess the risk of invasive species and for predicting species responses to climate change. Our findings challenge the tenet that species' niches are conserved aspects of their ecology.

Intraspecific diversity buffers the inhibitory effects of soil biota.

Plant community productivity can increase with increasing intraspecific genotypic diversity. Previous studies have attributed the genetic diversity-productivity pattern to differential resource use among genotypes, as many studies have found for species. But here we ask whether suppression of productivity at low intraspecific diversity by soil biota might also drive a positive diversity-productivity relationship. In a previous study, we manipulated genetic diversity by varying the number of Pseudoroegneria accessions growing together in common garden plots, and used soil from that experiment to evaluate soil feedbacks. The total biomass of P. spicata plants grown in unsterilized soil increased with accession richness, specifically when comparing soil that had contained plants from 3 accessions to soil that had contained plants from either 8 or 12 population accessions. Furthermore, soil from high-richness (8 or 12-accession) plots drove neutral feedbacks, whereas soil in the 3-accession plots (3) drove negative feedbacks. However, within each level of richness, there was no relationship between relative yield and feedback. Our results suggest that soil biota might play an integral role in the emerging understanding of the relationship between intraspecific diversity and ecosystem productivity.

Multi-Phase US Spread and Habitat Switching of a Post-Columbian Invasive, Sorghum halepense.

Johnsongrass (Sorghum halepense) is a striking example of a post-Columbian founder event. This natural experiment within ecological time-scales provides a unique opportunity for understanding patterns of continent-wide genetic diversity following range expansion. Microsatellite markers were used for population genetic analyses including leaf-optimized Neighbor-Joining tree, pairwise FST, mismatch analysis, principle coordinate analysis, Tajima's D, Fu's F and Bayesian clusterings of population structure. Evidence indicates two geographically distant introductions of divergent genotypes, which spread across much of the US in <200 years. Based on geophylogeny, gene flow patterns can be inferred to have involved five phases. Centers of genetic diversity have shifted from two introduction sites separated by ~2000 miles toward the middle of the range, consistent with admixture between genotypes from the respective introductions. Genotyping provides evidence for a 'habitat switch' from agricultural to non-agricultural systems and may contribute to both Johnsongrass ubiquity and aggressiveness. Despite lower and more structured diversity at the invasion front, Johnsongrass continues to advance northward into cooler and drier habitats. Association genetic approaches may permit identification of alleles contributing to the habitat switch or other traits important to weed/invasive management and/or crop improvement.

An exotic invasive plant selects for increased competitive tolerance, but not competitive suppression, in a native grass.

Exotic invasive plants can exert strong selective pressure for increased competitive ability in native plants. There are two fundamental components of competitive ability: suppression and tolerance, and the current paradigm that these components have equal influences on a species' overall competitive ability has been recently questioned. If these components do not have equal influences on overall ability, then selection on competitive tolerance and suppression may be disproportionate. We used naturally invaded communities to study the effects of selection caused by an invasive forb, Centaurea stoebe, on a native grass, Pseudoroegneria spicata. P. spicata plants were harvested from within dense C. stoebe patches and from nearby uninvaded areas, divided clonally into replicates, then transplanted into a common garden where they grew alone or competed with C. stoebe. We found that P. spicata plants collected from within C. stoebe patches were significantly more tolerant of competition with C. stoebe than P. spicata plants collected from uninvaded areas, but plants from inside invaded patches were not superior at suppressing C. stoebe. These results are consistent with the hypothesis that strong competitors may select for tolerance to competition more than for the ability to suppress neighbors. This has important implications for how native plant communities may respond to invasion over time, and how invasive and native species may ultimately coexist.

Root contact responses and the positive relationship between intraspecific diversity and ecosystem productivity.

High species and functional group richness often has positive effects on ecosystem function including increasing productivity. Recently, intraspecific diversity has been found to have similar effects, but because traits vary far less within a species than among species we have a much poorer understanding of the mechanisms by which intraspecific diversity affects ecosystem function. We explored the potential for identity recognition among the roots of different Pseudoroegneria spicata accessions to contribute to previously demonstrated overyielding in plots with high intraspecific richness of this species relative to monocultures. First, we found that when plants from different populations were planted together in pots the total biomass yield was 30 % more than in pots with two plants from the same population. Second, we found that the elongation rates of roots of Pseudoroegneria plants decreased more after contact with roots from another plant from the same population than after contact with roots from a plant from a different population. These results suggest the possibility of some form of detection and avoidance mechanism among more closely related Pseudoroegneria plants. If decreased growth after contact results in reduced root overlap, and reduced root overlap corresponds with reduced growth and productivity, then variation in detection and avoidance among related and unrelated accessions may contribute to how ecotypic diversity in Pseudoroegneria increases productivity.

Structural and functional characterization of the acidic region from the RIZ tumor suppressor.

RIZ (retinoblastoma protein-interacting zinc finger protein), also denoted PRDM2, is a transcriptional regulator and tumor suppressor. It was initially identified because of its ability to interact with another well-established tumor suppressor, the retinoblastoma protein (Rb). A short motif, IRCDE, in the acidic region (AR) of RIZ was reported to play an important role in the interaction with the pocket domain of Rb. The IRCDE motif is similar to a consensus Rb-binding sequence LXCXE (where X denotes any amino acid) that is found in several viral Rb-inactivating oncoproteins. To improve our understanding of the molecular basis of binding of Rb to RIZ, we investigated the interaction between purified recombinant AR and the pocket domain of Rb using nuclear magnetic resonance spectroscopy, isothermal titration calorimetry, and fluorescence anisotropy experiments. We show that AR is intrinsically disordered and that it binds the pocket domain with submicromolar affinity. We also demonstrate that the interaction between AR and the pocket domain is mediated primarily by the short stretch of residues containing the IRCDE motif and that the contribution of other parts of AR to the interaction with the pocket domain is minimal. Overall, our data provide clear evidence that RIZ is one of the few cellular proteins that can interact directly with the LXCXE-binding cleft on Rb.

Testing the mechanisms of diversity-dependent overyielding in a grass species.

Plant diversity enhances many ecosystem processes, including productivity, but these effects have been studied almost exclusively at the taxonomic scale of species. We explore the effect of intraspecific diversity on the productivity of a widespread and dominant grassland species using accessions collected from populations throughout its range. We found that increasing population/ecotype diversity of Pseudoroegneria spicata increased productivity to a similar degree as that reported for species diversity. However, we did not find evidence that overyielding was related to either resource depletion or to pathogenic soil fungi, two causes of overyielding in species-diverse communities. Instead, larger accessions overyielded at low diversity at the expense of smaller accessions, and small accessions overyielded through complementarity at all levels of diversity. Furthermore, overyielding was stronger for accessions from mesic environments, suggesting that local adaptation might predictably influence how plants respond to increases in diversity. This suggests that mass-based competition or other cryptic accession-specific processes had complex but important effects on overyielding. Our results indicate that the effects of diversity within a species can be substantial but that overyielding by intraspecifically diverse populations may not be through the same processes thought to cause overyielding in species diverse communities.

Inheritance, mode of inheritance, and candidate genes for primary hyperparathyroidism in Keeshonden.

BACKGROUND: Primary hyperparathyroidism (PHPT) is caused by inappropriate secretion of parathyroid hormone (PTH) by autonomously functioning neoplastic or hyperplastic parathyroid "chief" cells. Keeshonden are thought to be over-represented in studies on canine PHPT, but no proof of heritability or mode of inheritance has been published. The canine disease clinically resembles human familial isolated hyperparathyroidism (FIHP). HYPOTHESIS: Primary hyperparathyroidism in Keeshonden is genetically transmitted and is caused by a mutation in 1 of 4 genes implicated in human FIHP: MEN1, CASR, HRPT2, or RET. ANIMALS: Pedigrees consisting of 1647 Keeshonden were created including 219 Keeshonden with known PHPT phenotypes (69 positive). DNA samples were obtained from 176 of the 219 Keeshonden (34 positive). METHODS: Heritability and mode of inheritance were determined by segregation analysis. Canine homologs to the human genes were identified. Exons and surrounding intron regions were sequenced and scanned for sense-altering polymorphisms or polymorphisms that segregated with the disease. Messenger RNA from a parathyroid tumor of an affected Keeshond was analyzed for polymorphisms and splice alterations. RESULTS: PHPT follows an autosomal dominant mode of inheritance in Keeshonden with possible age-dependent penetrance. No polymorphisms identified in the genes analyzed were associated with a change in predicted protein or in hypothesized splice sites. CONCLUSIONS AND CLINICAL IMPORTANCE: PHPT is an autosomal dominant, genetically transmitted disease in Keeshonden. Once the mutation locus is identified, genetic testing should quickly decrease the incidence of PHPT in this breed. It is unlikely that mutations in MEN1, CASR, HRPT2, or RET cause PHPT in Keeshonden.