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Reliable Method for Assessing Seed Germination, Dormancy, and Mortality under Field Conditions

We describe techniques for approximating seed bank dynamics over time using Helianthus annuus as an example study species. Strips of permeable polyester ...

Ohio State University

Gene flow from crop fields to wild populations produces hybrids that often differ from their wild counterparts in growth form, phenology, and life history characteristics. Germination and dormancy dynamics have a strong influence on population persistence, competitive dynamics, and ultimately, plant fitness. They may also play a role in modifying crop gene introgression, which has been of primary interest since the release of transgenic crops. We investigated how seed germination and dormancy were affected by sunflower crop wild hybridization in both laboratory and field experiments. Hybridization increased seed germination and decreased dormancy. Of the nine wild populations we assayed, most of their hybrids had higher germination than the wilds of the same population. However, absolute germination levels varied by population and testing environment. Hybrids produced by three different crop lines differed in germination, and their germination rankings shifted across populations. Increased germination in hybrids could accelerate crop gene introgression, provided that hybrids germinate in an appropriate period. Differences in relative germination of wild and hybrid seed indicated that the effect of germination on introgression will likely vary by population due, in part, to initial levels of dormancy in the population. Therefore, the implications of gene flow from crops with novel characteristics or from transgenic crops will also vary by population.

Increased germination of diverse crop-wild hybrid sunflower seeds.

Gene flow between crop fields and wild populations often results in hybrids with reduced fitness compared to their wild counterparts due to characteristics imparted by the crop genome. But the specifics of the evolutionary outcome of crop-wild gene flow may depend on context, varying due to local environmental conditions and genetic variation within and among wild populations and among crop lines. To evaluate context-dependence of fitness of F1 hybrids, sunflower crop lines were crossed with nine wild populations from across the northern United States. These crop-wild hybrids and their wild counterparts were grown under agricultural conditions in the field with and without wheat competition. Hybrids were far less fecund than wild plants, yet more likely to survive to reproduce. There was considerable variability among wild populations for fecundity and the specific crop line used to generate the crop-wild hybrid significantly affected fecundity. The fitness deficit suffered by crop-wild hybrids varied by population, as did the rankings of the crop-wild hybrids from three different crop lines. Wheat competition decreased fecundity and survival considerably and hampered seed production of wild plants more than that of hybrids. Genotype x environment interactions indicated that the response of fitness to competition differed by population. Consequently, the fitness of hybrids relative to wild plants varied considerably among wild populations and was not consistent across environments. Notably, relative fitness of hybrids was greater under competitive conditions. This research is the first study of its kind to demonstrate that the consequences of crop-wild gene flow are context dependent and contingent on the genetics of the specific wild populations and the local biotic and abiotic conditions.

Effects of competition on the fitness of wild and crop-wild hybrid sunflower from a diversity of wild populations and crop lines.

After a decade of transgenic crop production, the dynamics of gene introgression into wild relatives remain unclear. Taking an ecological genetics approach to investigating fitness in crop-wild hybrid zones, we uncovered both conditions and characteristics that may promote introgression. We compared diverse crop-wild hybrid genotypes relative to wild Helianthus annuus under one benign and three stressful agricultural environments. Whereas relative fitness of crop-wild hybrids averaged 0.25 under benign conditions, with herbicide application or competition it reached 0.45 and was more variable. In some instances, hybrid fitness matched wild fitness (approximately 1). Thus, wild populations under agronomic stress may be more susceptible to introgression. Although 'domestication' traits are typically considered unlikely to persist in wild populations, we found some (e.g. rapid growth and early flowering) that may enhance hybrid fitness, especially in stressful environments. Rigorous assessment of how particular genotypes, phenotypes, and environments affect introgression will improve risk assessment for transgenic crops.

Stress and domestication traits increase the relative fitness of crop-wild hybrids in sunflower.

Adaptive nature of crop cytoplasm.

Variation in seedling emergence timing is considered adaptive over the long term in wild populations, but early emergence can result in a fitness advantage. To explore the adaptive significance of seedling emergence timing, it should be studied under realistic conditions and in the context of other traits that influence fitness.

Selection on seedling emergence timing and size in an annual plant, Helianthus annuus (common sunflower, Asteraceae).

• Premise of the study: The fitness of an offspring may depend on its nuclear genetic composition (via both parental genotypes) as well as on genetic maternal effects (via only the maternal parent). Understanding the relative importance of these two genetic factors is particularly important for research on crop-wild hybridization, since traits with important genetic maternal effects (e.g., seed size) often differ among crops and their relatives. We hypothesized that the effects of these genetic factors on fitness components would change across the life cycle of hybrids.• Methods: We followed seed, plant size, and reproductive traits in field experiments with wild and four crop-wild hybrids of sunflower (Helianthus annuus), which differed in nuclear genetic composition and maternal parent (wild or F1 hybrid).• Key results: We identified strong genetic maternal effects for early life cycle characteristics, with seeds produced on an F1 mother having premature germination, negligible seed dormancy, and greater seedling size. Increased percentages of crop alleles also increased premature germination and reduced dormancy in seeds produced on a wild mother. For mature plants, nuclear genetic composition dominated: greater percentages of crop alleles reduced height, branching, and fecundity.• Conclusions: Particular backcrosses between hybrids and wilds may differentially facilitate movement of crop alleles into wild populations due to their specific features. For example, backcross seeds produced on wild mothers can persist in the seed bank, illustrating the importance of genetic maternal effects, whereas backcross individuals with either wild or F1 mothers have high fecundity, resulting from their wild-like nuclear genetic composition.

Roles of maternal effects and nuclear genetic composition change across the life cycle of crop-wild hybrids.

Understanding the likelihood and extent of introgression of novel alleles in hybrid zones requires comparison of lifetime fitness of parents and hybrid progeny. However, fitness differences among cross types can vary depending on biotic conditions, thereby influencing introgression patterns. Based on past work, we predicted that increased competition would enhance introgression between cultivated and wild sunflower (Helianthus annuus) by reducing fitness advantages of wild plants. To test this prediction, we established a factorial field experiment in Kansas, USA where we monitored the fitness of four cross types (Wild, F1, F2, and BCw hybrids) under different levels of interspecific and intraspecific competition. Intraspecific manipulations consisted both of density of competitors and of frequency of crop-wild hybrids. We recorded emergence of overwintered seeds, survival to reproduction, and numbers of seeds produced per reproductive plant. We also calculated two compound fitness measures: seeds produced per emerged seedling and seeds produced per planted seed. Cross type and intraspecific competition affected emergence and survival to reproduction, respectively. Further, cross type interacted with competitive treatments to influence all other fitness traits. More intense competition treatments, especially related to density of intraspecific competitors, repeatedly reduced the fitness advantage of wild plants when considering seeds produced per reproductive plant and per emerged seedling, and F2 plants often became indistinguishable from the wilds. Wild fitness remained superior when seedling emergence was also considered as part of fitness, but the fitness of F2 hybrids relative to wild plants more than quadrupled with the addition of interspecific competitors and high densities of intraspecific competitors. Meanwhile, contrary to prediction, lower hybrid frequency reduced wild fitness advantage. These results emphasize the importance of taking a full life cycle perspective. Additionally, due to effects of exogenous selection, a given hybrid generation may be especially well-suited to hastening introgression under particular environmental conditions.

Fitness of crop-wild hybrid sunflower under competitive conditions: implications for crop-to-wild introgression.

Landraces cultivated in centers of crop diversity result from past and contemporary patterns of natural and farmer-mediated evolutionary forces. Successful in situ conservation of crop genetic resources depends on continuity of these evolutionary processes. Climate change is projected to affect agricultural production, yet analyses of impacts on in situ conservation of crop genetic diversity and farmers who conserve it have been absent. How will crop landraces respond to alterations in climate? We review the roles that phenotypic plasticity, evolution, and gene flow might play in sustaining production, although we might expect erosion of genetic diversity if landrace populations or entire races lose productivity. For example, highland maize landraces in southern Mexico do not express the plasticity necessary to sustain productivity under climate change, but may evolve in response to altered conditions. The outcome for any given crop in a given region will depend on the distribution of genetic variation that affects fitness and patterns of climate change. Understanding patterns of neutral and adaptive diversity from the population to the landscape scale is essential to clarify how landraces conserved in situ will continue to evolve and how to minimize genetic erosion of this essential natural resource.

Evolutionary response of landraces to climate change in centers of crop diversity.

Domestication has resulted in selection upon seed traits found in wild populations, yet crop-wild hybrids retain some aspects of both parental phenotypes. Seed fates of germination, dormancy, and mortality can influence the success of crop allele introgression in crop-wild hybrid zones, especially if crop alleles or crop-imparted seed coverings result in out-of-season germination. We performed a seed burial experiment using crop, wild, and diverse hybrid sunflower (Helianthus annuus) cross types to test how a cross type's maternal parent and nuclear genetic composition might affect its fate under field conditions. We observed higher maladaptive fall germination in the crop- and F1- produced seeds than wild-produced seeds and, due to an interaction with percent crop alleles, fall germination was higher for cross types with more crop-like nuclear genetics. By spring, crop-produced cross types had the highest overwintering mortality, primarily due to higher fall germination. Early spring germination was identical across maternal types, but germination continued for F1-produced seeds. In conclusion, the more wild-like the maternal parent or the less proportion of the cross type's genome contributed by the crop, the greater likelihood a seed will remain ungerminated than die. Wild-like dormancy may facilitate introgression through future recruitment from the soil seed bank.

Seed fates in crop-wild hybrid sunflower: crop allele and maternal effects.

Hybridization produces strong evolutionary forces. In hybrid zones, selection can differentially occur on traits and selection intensities may differ among hybrid generations. Understanding these dynamics in crop-wild hybrid zones can clarify crop-like traits likely to introgress into wild populations and the particular hybrid generations through which introgression proceeds. In a field experiment with four crop-wild hybrid Helianthus annuus (sunflower) cross types, we measured growth and life history traits and performed phenotypic selection analysis on early season traits to ascertain the likelihood, and routes, of crop allele introgression into wild sunflower populations. All cross types overwintered, emerged in the spring, and survived until flowering, indicating no early life history barriers to crop allele introgression. While selection indirectly favored earlier seedling emergence and taller early season seedlings, direct selection only favored greater early season leaf length. Further, there was cross type variation in the intensity of selection operating on leaf length. Thus, introgression of multiple early season crop-like traits, due to direct selection for greater early season leaf length, should not be impeded by any cross type and may proceed at different rates among generations. In sum, alleles underlying early season sunflower crop-like traits are likely to introgress into wild sunflower populations.

Kristin L. Mercer

Department of Horticulture and Crop Science

Ohio State University

Reliable Method for Assessing Seed Germination, Dormancy, and Mortality under Field Conditions