Discussion: Prioritize perennial grain development for sustainable food production and environmental benefits.

Perennial grains have potential to contribute to ecological intensification of food production by enabling the direct harvest of human-edible crops without requiring annual cycles of disturbance and replanting. Studies of prototype perennial grains and other herbaceous perennials point to the ability of agroecosystems including these crops to protect water quality, enhance wildlife habitat, build soil quality, and sequester soil carbon. However, genetic improvement of perennial grain candidates has been hindered by limited investment due to uncertainty about whether the approach is viable. As efforts to develop perennial grain crops have expanded in past decades, critiques of the approach have arisen. With a recent report of perennial rice producing yields equivalent to those of annual rice over eight consecutive harvests, many theoretical concerns have been alleviated. Some valid questions remain over the timeline for new crop development, but we argue these may be mitigated by implementation of recent technological advances in crop breeding and genetics such as low-cost genotyping, genomic selection, and genome editing. With aggressive research investment in the development of new perennial grain crops, they can be developed and deployed to provide atmospheric greenhouse gas reductions.

Nitrate leaching losses and the fate of 15N fertilizer in perennial intermediate wheatgrass and annual wheat - A field study.

Perennial grains, such as the intermediate wheatgrass (Thinopyrum intermedium) (IWG), may reduce negative environmental effects compared to annual grain crops. Their permanent, and generally larger, root systems are likely to retain nitrogen (N) better, decreasing harmful losses of N and improving fertilizer N use efficiency, but there have been no comprehensive N fertilizer recovery studies in IWG to date. We measured fertilizer N recovery with stable isotope tracers in crop biomass and soil, soil N mineralization and nitrification, and nitrate leaching in IWG and annual wheat in a replicated block field experiment. Nitrate leaching was drastically reduced in IWG (0.1 and 3.1 kg N ha-1 yr-1) in its third and fourth year since establishment, compared with 5.6 kg N ha-1 yr-1 in annual wheat and 41.0 kg N ha-1 yr-1 in fallow respectively. There were no differences in net N mineralization or nitrification between IWG and annual wheat, though there was generally more inorganic N in the soil profile of annual wheat. More 15N fertilizer was recovered in the straw and all depths of the roots and soils in IWG than annual wheat. However, annual wheat recovered much more 15N fertilizer in the seeds compared to IWG, which had lower grain yields. 15N-labeled fertilizer contributed little (<3 %) to nitrate-N in leachate, highlighting the role of soil microbes in regulating loss of current year fertilizer N. The large reduction in nitrate leaching demonstrates that perennial grains can reduce harmful nitrogen losses and offer a more sustainable alternative to annual grains.

Perennials as Future Grain Crops: Opportunities and Challenges.

Perennial grain crops could make a valuable addition to sustainable agriculture, potentially even as an alternative to their annual counterparts. The ability of perennials to grow year after year significantly reduces the number of agricultural inputs required, in terms of both planting and weed control, while reduced tillage improves soil health and on-farm biodiversity. Presently, perennial grain crops are not grown at large scale, mainly due to their early stages of domestication and current low yields. Narrowing the yield gap between perennial and annual grain crops will depend on characterizing differences in their life cycles, resource allocation, and reproductive strategies and understanding the trade-offs between annualism, perennialism, and yield. The genetic and biochemical pathways controlling plant growth, physiology, and senescence should be analyzed in perennial crop plants. This information could then be used to facilitate tailored genetic improvement of selected perennial grain crops to improve agronomic traits and enhance yield, while maintaining the benefits associated with perennialism.

Carbon and water relations in perennial Kernza (Thinopyrum intermedium): An overview.

Perennial crops have been proposed as a more sustainable alternative to annual crops, because they have extended growing seasons, continuous ground cover, reduced nutrient leakage, and sequester more carbon in the soils than annual crops. One example is intermediate wheatgrass (Thinopyrum intermedium), a perennial crop that has been used as a cool-season forage throughout the USA and Canada and also across its native range in Eurasia. Since the 1980's, intermediate wheatgrass has been under domestication to improve seed fertility and grain yield. Commercial products are being sold under the trade name Kernza, owned by The Land Institute, located in Salina, Kansas, USA. This review provides a comprehensive framework about the physical and biological aspects involving the water and carbon cycles in Kernza plants. The main aspects we highlight here are based on previous findings regarding Kernza: i) the ability of maintaining a relatively high water-use efficiency throughout the whole growing season, which is beneficial to mitigate water stress, representing an important physiological mean to acclimate under severe, unfavorable weather conditions, and ii) its higher evapotranspiration (ET) and net carbon uptake rates, particularly when compared to annual counterparts. Only a thorough multifaceted assessment of the repercussion for carbon and water fluxes of a shift from annual crops to Kernza will allow assessing the perspectives of such novel perennial crop to support food security and a number of ecosystem services, particularly under future climates.

New Food Crop Domestication in the Age of Gene Editing: Genetic, Agronomic and Cultural Change Remain Co-evolutionarily Entangled.

The classic domestication scenario for grains and fruits has been portrayed as the lucky fixation of major-effect "domestication genes." Characterization of these genes plus recent improvements in generating novel alleles (e.g., by gene editing) have created great interest in de novo domestication of new crops from wild species. While new gene editing technologies may accelerate some genetic aspects of domestication, we caution that de novo domestication should be understood as an iterative process rather than a singular event. Changes in human social preferences and relationships and ongoing agronomic innovation, along with broad genetic changes, may be foundational. Allele frequency changes at many loci controlling quantitative traits not normally included in the domestication syndrome may be required to achieve sufficient yield, quality, defense, and broad adaptation. The environments, practices and tools developed and maintained by farmers and researchers over generations contribute to crop yield and success, yet those may not be appropriate for new crops without a history of agronomy. New crops must compete with crops that benefit from long-standing participation in human cultural evolution; adoption of new crops may require accelerating the evolution of new crops' culinary and cultural significance, the emergence of markets and trade, and the formation and support of agricultural and scholarly institutions. We provide a practical framework that highlights and integrates these genetic, agronomic, and cultural drivers of change to conceptualize de novo domestication for communities of new crop domesticators, growers and consumers. Major gene-focused domestication may be valuable in creating allele variants that are critical to domestication but will not alone result in widespread and ongoing cultivation of new crops. Gene editing does not bypass or diminish the need for classical breeding, ethnobotanical and horticultural knowledge, local agronomy and crop protection research and extension, farmer participation, and social and cultural research and outreach. To realize the ecological and social benefits that a new era of de novo domestication could offer, we call on funding agencies, proposal reviewers and authors, and research communities to value and support these disciplines and approaches as essential to the success of the breakthroughs that are expected from gene editing techniques.

Abiotic and biotic context dependency of perennial crop yield.

Perennial crops in agricultural systems can increase sustainability and the magnitude of ecosystem services, but yield may depend upon biotic context, including soil mutualists, pathogens and cropping diversity. These biotic factors themselves may interact with abiotic factors such as drought. We tested whether perennial crop yield depended on soil microbes, water availability and crop diversity by testing monocultures and mixtures of three perennial crop species: a novel perennial grain (intermediate wheatgrass-Thinopyrum intermedium-- that produces the perennial grain Kernza®), a potential perennial oilseed crop (Silphium intregrifolium), and alfalfa (Medicago sativa). Perennial crop performance depended upon both water regime and the presence of living soil, most likely the arbuscular mycorrhizal (AM) fungi in the whole soil inoculum from a long term perennial monoculture and from an undisturbed native remnant prairie. Specifically, both Silphium and alfalfa strongly benefited from AM fungi. The presence of native prairie AM fungi had a greater benefit to Silphium in dry pots and alfalfa in wet pots than AM fungi present in the perennial monoculture soil. Kernza did not benefit from AM fungi. Crop mixtures that included Kernza overyielded, but overyielding depended upon inoculation. Specifically, mixtures with Kernza overyielded most strongly in sterile soil as Kernza compensated for poor growth of Silphium and alfalfa. This study identifies the importance of soil biota and the context dependence of benefits of native microbes and the overyielding of mixtures in perennial crops.

Comparative Analysis of Early Life Stage Traits in Annual and Perennial Phaseolus Crops and Their Wild Relatives.

Herbaceous perennial species are receiving increased attention for their potential to provide both edible products and ecosystem services in agricultural systems. Many legumes (Fabaceae Lindl.) are of special interest due to nitrogen fixation carried out by bacteria in their roots and their production of protein-rich, edible seeds. However, herbaceous perennial legumes have yet to enter widespread use as pulse crops, and the response of wild, herbaceous perennial species to artificial selection for increased seed yield remains under investigation. Here we compare cultivated and wild accessions of congeneric annual and herbaceous perennial legume species to investigate associations of lifespan and cultivation with early life stage traits including seed size, germination, and first year vegetative growth patterns, and to assess variation and covariation in these traits. We use "cultivated" to describe accessions with a history of human planting and use, which encompasses a continuum of domestication. Analyses focused on three annual and four perennial species of the economically important genus Phaseolus. We found a significant association of both lifespan and cultivation status with seed size (weight, two-dimensional lateral area, length), node number, and most biomass traits (with cultivation alone showing additional significant associations). Wild annual and perennial accessions primarily showed only slight differences in trait values. Relative to wild forms, both cultivated annual and cultivated perennial accessions exhibited greater seed size and larger overall vegetative size, with cultivated perennials showing greater mean trait differences relative to wild accessions than cultivated annuals. Germination proportion was significantly lower in cultivated relative to wild annual accessions, while no significant difference was observed between cultivated and wild perennial germination. Regardless of lifespan and cultivation status, seed size traits were positively correlated with most vegetative traits, and all biomass traits examined here were positively correlated. This study highlights some fundamental similarities and differences between annual and herbaceous perennial legumes and provides insights into how perennial legumes might respond to artificial selection compared to annual species.

Digging Deeper for Agricultural Resources, the Value of Deep Rooting.

In the quest for sustainable intensification of crop production, we discuss the option of extending the root depth of crops to increase the volume of soil exploited by their root systems. We discuss the evidence that deeper rooting can be obtained by appropriate choice of crop species, by plant breeding, or crop management and its potential contributions to production and sustainable development goals. Many studies highlight the potentials of deeper rooting, but we evaluate its contributions to sustainable intensification of crop production, the causes of the limited research into deep rooting of crops, and the research priorities to fill the knowledge gaps.

Community structure of soil fungi in a novel perennial crop monoculture, annual agriculture, and native prairie reconstruction.

The use of perennial crop species in agricultural systems may increase ecosystem services and sustainability. Because soil microbial communities play a major role in many processes on which ecosystem services and sustainability depend, characterization of soil community structure in novel perennial crop systems is necessary to understand potential shifts in function and crop responses. Here, we characterized soil fungal community composition at two depths (0-10 and 10-30 cm) in replicated, long-term plots containing one of three different cropping systems: a tilled three-crop rotation of annual crops, a novel perennial crop monoculture (Intermediate wheatgrass, which produces the grain Kernza®), and a native prairie reconstruction. The overall fungal community was similar under the perennial monoculture and native vegetation, but both were distinct from those in annual agriculture. The mutualist and saprotrophic community subsets mirrored differences of the overall community, but pathogens were similar among cropping systems. Depth structured overall communities as well as each functional group subset. These results reinforce studies showing strong effects of tillage and sampling depth on soil community structure and suggest plant species diversity may play a weaker role. Similarities in the overall and functional fungal communities between the perennial monoculture and native vegetation suggest Kernza® cropping systems have the potential to mimic reconstructed natural systems.

Drought exacerbates negative consequences of high-intensity cattle grazing in a semiarid grassland.

Grasslands managed for grazing are the largest land-use category globally, with a significant proportion of these grasslands occurring in semiarid and arid regions. In such dryland systems, the effect of grazing on native plant diversity has been equivocal, some studies suggesting that grazing reduces native plant diversity, others that grazing increases or has little impact on diversity. One impediment toward generalizing grazing effects on diversity in this region is that high levels of interannual variation in precipitation may obfuscate vegetative response patterns. By analyzing a long-term data set collected over a 20-yr period in a semiarid grassland, we explicitly evaluated the role of climate in regulating the effect of cattle grazing on plant communities, finding that climate interacted with grazing intensity to shape grassland communities. Community composition of plots that were intensively grazed varied considerably in response to climatic variation and native species richness was low relative to ungrazed and moderately grazed plots. Following a severe drought in 2002, exotic species richness rapidly increased in the high-intensity grazing plots. While this pattern was mirrored in the other treatments, exotic species richness increased to a greater extent and was slower to return to pre-drought levels in the high-intensity grazing plots. Overall, moderate grazing, even compared to grazing cessation, stabilized grassland communities through time, increased resilience to drought, and maintained the highest levels of native plant diversity and lowest levels of exotic diversity. These findings suggest that grazing, at moderate levels, may support grassland resilience to climate change in semiarid regions. However, grazing that exceeds tolerances, particularly in combination with extreme climatic events, like drought, can alter plant composition over relatively long timescales and possibly increase invasibility by nonnative species.

Managing for Multifunctionality in Perennial Grain Crops.

Plant breeders are increasing yields and improving agronomic traits in several perennial grain crops, the first of which is now being incorporated into commercial food products. Integration strategies and management guidelines are needed to optimize production of these new crops, which differ substantially from both annual grain crops and perennial forages. To offset relatively low grain yields, perennial grain cropping systems should be multifunctional. Growing perennial grains for several years to regenerate soil health before rotating to annual crops and growing perennial grains on sloped land and ecologically sensitive areas to reduce soil erosion and nutrient losses are two strategies that can provide ecosystem services and support multifunctionality. Several perennial cereals can be used to produce both grain and forage, and these dual-purpose crops can be intercropped with legumes for additional benefits. Highly diverse perennial grain polycultures can further enhance ecosystem services, but increased management complexity might limit their adoption.

Can evolutionary constraints explain the rarity of nitrogen-fixing trees in high-latitude forests?

Contents 1195 I. 1195 II. 1196 III. 1196 IV. 1200 1200 References 1200 SUMMARY: The rarity of symbiotic nitrogen (N)-fixing trees in temperate and boreal ('high-latitude') forests is curious. One explanation - the evolutionary constraints hypothesis - posits that high-latitude N-fixing trees are rare because few have evolved. Here, we consider traits necessary for high-latitude N-fixing trees. We then use recent developments in trait evolution to estimate that > 2000 and > 500 species could have evolved from low-latitude N-fixing trees and high-latitude N-fixing herbs, respectively. Evolution of N-fixing from nonfixing trees is an unlikely source of diversity. Dispersal limitation seems unlikely to limit high-latitude N-fixer diversity. The greater number of N-fixing species predicted to evolve than currently inhabit high-latitude forests suggests a greater role for ecological than evolutionary constraints.

Impact of grazing intensity during drought in an Arizona grassland.

The ecological benefits of changing cattle grazing practices in the western United States remain controversial, due in part to a lack of experimentation. In 1997 we initiated an experimental study of two rangeland alternatives, cattle removal and high-impact grazing, and compared grassland community responses with those with more conventional, moderate grazing practices. The study was conducted in a high-elevation, semiarid grassland near Flagstaff, Arizona (U.S.A.). We conducted annual plant surveys of modified Whittaker plots for 8 years and examined plant composition shifts among treatments and years. High-impact grazing had strong directional effects that led to a decline in perennial forb cover and an increase in annual plants, particularly the exotic cheatgrass (Bromus tectorum L.). A twofold increase in plant cover by exotic species followed a severe drought in the sixth year of the study, and this increase was greatest in the high-impact grazing plots, where native cover declined by one-half. Cattle removal resulted in little increase in native plant cover and reduced plant species richness relative to the moderate grazing control. Our results suggest that some intermediate level of cattle grazing may maintain greater levels of native plant diversity than the alternatives of cattle removal or high-density, short-duration grazing practices. Furthermore, episodic drought interacts with cattle grazing, leading to infrequent, but biologically important shifts in plant communities. Our results demonstrate the importance of climatic variation in determining ecological effects of grazing practices, and we recommend improving conservation efforts in arid rangelands by developing management plans that anticipate this variation.

Self-efficacy and pain behavior among subjects with fibromyalgia.

Given the lack of objective physical measures for assessing fibromyalgia syndrome (FS), the role of pain assessment is particularly important. The role of psychological factors is controversial among FS patients. This study was designed to better understand the relationship between pain behaviors and psychological variables. Specifically, this study (1) refined a pain behavior observation (PBO) methodology for use with FS patients, (2) determined whether stretching is a valid pain behavior, and (3) assessed whether psychological variables including self-efficacy and/or depression can predict pain behaviors after controlling for disease severity and age. The 73 FS subjects meeting the American College of Rheumatology classification system completed questionnaires measuring self-efficacy, depression, and pain. Trained physicians conducted tender-point examinations. Subjects were video-taped using a standardized procedure. Two trained raters independently coded all pain behaviors. Kappa coefficients and correlations among pain behaviors and self-reported pain indicated that the PBO method was both reliable and valid. However, the newly defined pain behavior 'stretching' was found to be negatively associated with self-reported pain. Hierarchical multiple regression (MR) analyses revealed that depression did not predict pain behavior over and above myalgic scores and age; however, in 3 separate MR analyses, self-efficacy for function, pain, and other symptoms each predicted pain behavior over and above myalgic scores and age. This study indicated that the original pain behavior scoring methodology is appropriate for use with the FS population and should not be modified to include the pain behavior 'stretching'. Self-efficacy was related to pain behavior while depression was not among this FS sample.

Pain behavior predictors among fibromyalgia patients.

This study was designed to assess the reliability and validity of a Pain Behavior Observation method with fibromyalgia syndrome (FS) subjects and to determine the factors which predict pain behavior among FS subjects. Fifty-eight female FS subjects participated in the videotaped Pain Behavior Observation method. Subjects also completed the McGill Pain Questionnaire (MPQ), the Arthritis Impact Measurement Scale (AIMS), and the Symptoms Checklist-90-Revised (SCL-90-R). Pain behavior totals were derived using the Pain Behavior Observation method of McDaniel et al. (1986). Results indicated that the scoring method of Keefe and Block (1982) developed for use with back pain patients was more generalizable to this FS sample. Total pain behavior score using the Keefe and Block scoring method was correlated with self-report pain. Interrater reliability ranged from 0.9 to 0.99. Hierarchical multiple regression analyses revealed that age and physical disability predicted the total pain behaviors. The Pain Behavior Observation method appears a valid pain measurement technique independent of depression among FS patients. Directions for future research are discussed.