Seeing the fields through the weeds: introducing the WeedEco R package for comparing past and present arable farming systems using functional weed ecology.

The functional ecology of arable weeds provides a way of comparing present-day and past farming regimes. This paper presents the R package WeedEco, an open-source resource which allows users to compare their archaeobotanical dataset against three previously published arable weed models to understand fertility, disturbance or a combination of both. The package provides functions for data organisation, classification and visualisation, allowing users to enter raw archaeobotanical data, obtain trait values from the functional trait dataset, conduct discriminant analysis and plot the results against the relevant present-day model. Using data from the early medieval site of Stafford in the UK, the paper provides a detailed example of the use of the package, demonstrating its different functions, as well as how the results can be interpreted. Supplementary Information: The online version contains supplementary material available at 10.1007/s00334-023-00964-8.

Genetic diversity, distribution and domestication history of the neglected GGAtAt genepool of wheat.

KEY MESSAGE: We present a comprehensive survey of cytogenetic and genomic diversity of the GGAtAt genepool of wheat, thereby unlocking these plant genetic resources for wheat improvement. Wheat yields are stagnating around the world and new sources of genes for resistance or tolerances to abiotic traits are required. In this context, the tetraploid wheat wild relatives are among the key candidates for wheat improvement. Despite its potential huge value for wheat breeding, the tetraploid GGAtAt genepool is largely neglected. Understanding the population structure, native distribution range, intraspecific variation of the entire tetraploid GGAtAt genepool and its domestication history would further its use for wheat improvement. The paper provides the first comprehensive survey of genomic and cytogenetic diversity sampling the full breadth and depth of the tetraploid GGAtAt genepool. According to the results obtained, the extant GGAtAt genepool consists of three distinct lineages. We provide detailed insights into the cytogenetic composition of GGAtAt wheats, revealed group- and population-specific markers and show that chromosomal rearrangements play an important role in intraspecific diversity of T. araraticum. The origin and domestication history of the GGAtAt lineages is discussed in the context of state-of-the-art archaeobotanical finds. We shed new light on the complex evolutionary history of the GGAtAt wheat genepool and provide the basis for an increased use of the GGAtAt wheat genepool for wheat improvement. The findings have implications for our understanding of the origins of agriculture in southwest Asia.

Fertile Crescent crop progenitors gained a competitive advantage from large seedlings.

Cereal domestication during the transition to agriculture resulted in widespread food production, but why only certain species were domesticated remains unknown. We tested whether seedlings of crop progenitors share functional traits that could give them a competitive advantage within anthropogenic environments, including higher germination, greater seedling survival, faster growth rates, and greater competitive ability.Fifteen wild grass species from the Fertile Crescent were grown individually under controlled conditions to evaluate differences in growth between cereal crop progenitors and other wild species that were never domesticated. Differences in germination, seedling survival, and competitive ability were measured by growing a subset of these species as monocultures and mixtures.Crop progenitors had greater germination success, germinated more quickly and had greater aboveground biomass when grown in competition with other species. There was no evidence of a difference in seedling survival, but seed size was positively correlated with a number of traits, including net assimilation rates, greater germination success, and faster germination under competition. In mixtures, the positive effect of seed mass on germination success and speed of germination was even more beneficial for crop progenitors than for other wild species, suggesting greater fitness. Thus, selection for larger seeded individuals under competition may have been stronger in the crop progenitors.The strong competitive ability of Fertile Crescent cereal crop progenitors, linked to their larger seedling size, represents an important ecological difference between these species and other wild grasses in the region. It is consistent with the hypothesis that competition within plant communities surrounding human settlements, or under early cultivation, benefited progenitor species, favoring their success as crops.

Seed size, number and strategies in annual plants: a comparative functional analysis and synthesis.

BACKGROUND AND AIMS: Plants depend fundamentally on establishment from seed. However, protocols in trait-based ecology currently estimate seed size but not seed number. This can be rectified. For annuals, seed number should simply be a positive function of vegetative biomass and a negative function of seed size. METHODS: Using published values of comparative seed number as the 'gold standard' and a large functional database, comparative seed yield and number per plant and per m2 were predicted by multiple regression. Subsequently, ecological variation in each was explored for English and Spanish habitats, newly calculated C-S-R strategies and changed abundance in the British flora. KEY RESULTS: As predicted, comparative seed mass yield per plant was consistently a positive function of plant size and competitive ability, and largely independent of seed size. Regressions estimating comparative seed number included, additionally, seed size as a negative function. Relationships differed numerically between regions, habitats and C-S-R strategies. Moreover, some species differed in life history over their geographical range. Comparative seed yield per m2 was positively correlated with FAO crop yield, and increasing British annuals produced numerous seeds. Nevertheless, predicted values must be viewed as comparative rather than absolute: they varied according to the 'gold standard' predictor used. Moreover, regressions estimating comparative seed yield per m2 achieved low precision. CONCLUSIONS: For the first time, estimates of comparative seed yield and number for >800 annuals and their predictor equations have been produced and the ecological importance of these regenerative traits has been illustrated. 'Regenerative trait-based ecology' remains in its infancy, with work needed on determinate vs. indeterminate flowering ('bet-hedging'), C-S-R methodologies, phylogeny, comparative seed yield per m2 and changing life history. Nevertheless, this has been a positive start and readers are invited to use estimates for >800 annuals, in the Supplementary data, to help advance 'regenerative trait-based ecology' to the next level.

Diversity of a cytokinin dehydrogenase gene in wild and cultivated barley.

The cytokinin dehydrogenase gene HvCKX2.1 is the regulatory target for the most abundant heterochromatic small RNAs in drought-stressed barley caryopses. We investigated the diversity of HvCKX2.1 in 228 barley landraces and 216 wild accessions and identified 14 haplotypes, five of these with ten or more members, coding for four different protein variants. The third largest haplotype was abundant in wild accessions (51 members), but absent from the landrace collection. Protein structure predictions indicated that the amino acid substitution specific to haplotype 3 could result in a change in the functional properties of the HvCKX2.1 protein. Haplotypes 1-3 have overlapping geographical distributions in the wild population, but the average rainfall amounts at the collection sites for haplotype 3 plants are significantly higher during November to February compared to the equivalent data for plants of haplotypes 1 and 2. We argue that the likelihood that haplotype 3 plants were excluded from landraces by sampling bias that occurred when the first wild barley plants were taken into cultivation is low, and that it is reasonable to suggest that plants with haplotype 3 are absent from the crop because these plants were less suited to the artificial conditions associated with cultivation. Although the cytokinin signalling pathway influences many aspects of plant development, the identified role of HvCKX2.1 in the drought response raises the possibility that the particular aspect of cultivation that mitigated against haplotype 3 relates in some way to water utilization. Our results therefore highlight the possibility that water utilization properties should be looked on as a possible component of the suite of physiological adaptations accompanying the domestication and subsequent evolution of cultivated barley.

Re-analysis of archaeobotanical remains from pre- and early agricultural sites provides no evidence for a narrowing of the wild plant food spectrum during the origins of agriculture in southwest Asia.

Archaeobotanical evidence from southwest Asia is often interpreted as showing that the spectrum of wild plant foods narrowed during the origins of agriculture, but it has long been acknowledged that the recognition of wild plants as foods is problematic. Here, we systematically combine compositional and contextual evidence to recognise the wild plants for which there is strong evidence of their deliberate collection as food at pre-agricultural and early agricultural sites across southwest Asia. Through sample-by-sample analysis of archaeobotanical remains, a robust link is established between the archaeological evidence and its interpretation in terms of food use, which permits a re-evaluation of the evidence for the exploitation of a broad spectrum of wild plant foods at pre-agricultural sites, and the extent to which this changed during the development of early agriculture. Our results show that relatively few of the wild taxa found at pre- and early agricultural sites can be confidently recognised as contributing to the human diet, and we found no evidence for a narrowing of the plant food spectrum during the adoption of agriculture. This has implications for how we understand the processes leading to the domestication of crops, and points towards a mutualistic relationship between people and plants as a driving force during the development of agriculture.

Diversity of a wall-associated kinase gene in wild and cultivated barley.

Domestication of barley and other cereals was accompanied by an increase in seed size which has been ascribed to human selection, large seeds being preferred by early farmers or favoured by cultivation practices such as deep sowing. An alternative suggestion is that the increase in seed size was an indirect consequence of selection for plants with more vigorous growth. To begin to address the latter hypothesis we studied the diversity of HvWAK1, a wall-associated kinase gene involved in root proliferation, in 220 wild barley accessions and 200 domesticated landraces. A 3655-bp sequence comprising the gene and upstream region contained 69 single nucleotide polymorphisms (SNPs), one indel and four short tandem repeats. A network of 50 haplotypes revealed a complex evolutionary relationship, but with landraces largely restricted to two parts of the topology. SNPs in the HvWAK1 coding region resulted in nonsynonymous substitutions at nine positions in the translation product, but none of these changes were predicted to have a significant effect on the protein structure. In contrast, the region upstream of the coding sequence contained five SNPs that were invariant in the domesticated population, fixation of these SNPs decreasing the likelihood that the upstream of a pair of TATA boxes and transcription start sites would be used to promote transcription of HvWAK1. The sequence diversity therefore suggests that the cis-regulatory region of HvWAK1 might have been subject to selection during barley domestication. The extent of root proliferation has been linked with traits such as above-ground biomass, so selection for particular cis-regulatory variants of HvWAK1 would be consistent with the hypothesis that seed size increases during domestication were the indirect consequence of selection for plants with increased growth vigour.

Phylogenetic patterns and phenotypic profiles of the species of plants and mammals farmed for food.

The origins of agriculture were key events in human history, during which people came to depend for their food on small numbers of animal and plant species. However, the biological traits determining which species were domesticated for food provision, and which were not, are unclear. Here, we investigate the phylogenetic distribution of livestock and crops, and compare their phenotypic traits with those of wild species. Our results indicate that phylogenetic clustering is modest for crop species but more intense for livestock. Domesticated species explore a reduced portion of the phenotypic space occupied by their wild counterparts and have particular traits in common. For example, herbaceous crops are globally characterized by traits including high leaf nitrogen concentration and tall canopies, which make them fast-growing species and proficient competitors. Livestock species are relatively large mammals with low basal metabolic rates, which indicate moderate to slow life histories. Our study therefore reveals ecological differences in domestication potential between plants and mammals. Domesticated plants belong to clades with traits that are advantageous in intensively managed high-resource habitats, whereas domesticated mammals are from clades adapted to moderately productive environments. Combining comparative phylogenetic methods with ecologically relevant traits has proven useful to unravel the causes and consequences of domestication.

Cereal progenitors differ in stand harvest characteristics from related wild grasses.

The domestication of crops in the Fertile Crescent began approximately 10,000 years ago indicating a change from a hunter-gatherer lifestyle to a sedentary, agriculture-based existence. The exploitation of wild plants changed during this transition, such that a small number of crops were domesticated from the broader range of species gathered from the wild. However, the reasons for this change are unclear.Previous studies have shown unexpectedly that crop progenitors are not consistently higher yielding than related wild grass species, when growing without competition. In this study, we replicate more closely natural competition within wild stands, using two greenhouse experiments to investigate whether cereal progenitors exhibit a greater seed yield per unit area than related wild species that were not domesticated.Stands of cereal progenitors do not provide a greater total seed yield per unit ground area than related wild species, but these crop progenitors do have greater reproductive efficiency than closely related wild species, with nearly twice the harvest index (the ratio of harvested seeds to total shoot dry mass).These differences arise because the progenitors have greater seed yield per tiller than closely related wild species, due to larger individual seed size but no reduction in seed number per tiller. The harvest characteristics of cereal progenitors may have made them a more attractive prospect than closely related wild species for the early cultivators who first planted these species, or could suggest an ecological filtering mechanism. Synthesis. Overall, we show that the maintenance of a high harvest index under competition, the packaging of seed in large tillers, and large seeds, consistently distinguish crop progenitors from closely related wild grass species. However, the archaeological significance of these findings remains unclear, since a number of more distantly related species, including wild oats, have an equally high or higher harvest index and yield than some of the progenitor species. Domestication of the earliest cereal crops from the pool of wild species available cannot therefore be explained solely by species differences in yield and harvest characteristics, and must also consider other plant traits.