Lessons from the history of Agave: ecological and cultural context for valuation of CAM.

BACKGROUND AND SCOPE: Crassulacean acid metabolism (CAM) is an intriguing physiological adaptation in plants that are widespread throughout many ecosystems. Despite the relatively recent mechanistic understanding of CAM in plant physiology, evidence from historical records suggests that ancient cultures in the Americas also recognized the value of CAM plants. Agave species, in particular, have a rich cultural legacy that provides a foundation for commercially valued products. Here, we review that legacy and potential relationships between ancient values and the needs of modern-day climate adaptation strategies. CONCLUSIONS: There are many products that can be produced from Agave species, including food, sugar, fibre and medicines. Traditional knowledge about agricultural management and preparation of plant products can be combined with new ecophysiological knowledge and agronomic techniques to develop these resources in the borderland region of the southwestern USA and Mexico. Historical records of pre-Columbian practices in the Sonoran desert and remnants of centuries-old agriculture in Baja California and Sonora demonstrate the climate resilience of Agave agriculture. Commercial growth of both tequila and bacanora indicates the potential for large-scale production today, but also underscores the importance of adopting regenerative agricultural practices to accomplish environmentally sustainable production. Recent international recognition of the Appellation of Origin for several Agave species produced for spirits in Mexico might provide opportunities for agricultural diversification. In contrast, fibre is currently produced from several Agave species on many continents. Projections of growth with future climate change suggest that Agave spp. will be viable alternatives for commodity crops that suffer declines during drought and increased temperatures. Historical cultivation of Agave affirms that these CAM plants can supply sugar, soft and hard fibres, medicines and food supplements.

A model of environmental limitations on production of Agave americana L. grown as a biofuel crop in semi-arid regions.

Plants that use crassulacean acid metabolism (CAM) have the potential to meet growing agricultural resource demands using land that is considered unsuitable for many common crop species. Agave americana L., an obligate CAM plant, has potential as an advanced biofuel crop in water-limited regions, and has greater cold tolerance than other high-yielding CAM species, but physiological tolerances have not been completely resolved. We developed a model to estimate the growth responses of A. americana to water input, temperature, and photosynthetically active radiation (PAR). The photosynthetic response to PAR was determined experimentally by measuring the integrated leaf gas exchange over 24 h after acclimation to six light levels. Maximum CO2 fixation rates were observed at a PAR intensity of 1250 µmol photons m-2 s-1. Growth responses of A. americana to water and temperature were also determined, and a monthly environmental productivity index (EPI) was derived that can be used to predict biomass growth. The EPI was calculated as the product of water, temperature, and light indices estimated for conditions at a site in Maricopa (Arizona), and compared with measured biomass at the same site (where the first field trial of A. americana as a crop was completed). The monthly EPI summed over the lifetime of multi-year crops was highly correlated with the average measured biomass of healthy 2- and 3-year-old plants grown in the field. The resulting relationship between EPI and biomass provides a simple model for estimating the production of A. americana at a monthly time step according to light, temperature, and precipitation inputs, and is a useful tool for projecting the potential geographic range of this obligate CAM species in future climatic conditions.

Undervalued potential of crassulacean acid metabolism for current and future agricultural production.

The potential for crassulacean acid metabolism (CAM) to support resilient crops that meet demands for food, fiber, fuel, and pharmaceutical products far exceeds current production levels. This review provides background on five families of plants that express CAM, including examples of many species within these families that have potential agricultural uses. We summarize traditional uses, current developments, management practices, environmental tolerance ranges, and economic values of CAM species with potential commercial applications. The primary benefit of CAM in agriculture is high water use efficiency that allows for reliable crop yields even in drought conditions. Agave species, for example, grow in arid conditions and have been exploited for agricultural products in North and South America for centuries. Yet, there has been very little investment in agricultural improvement for most useful Agave varieties. Other CAM species that are already traded globally include Ananas comosus (pineapple), Aloe spp., Vanilla spp., and Opuntia spp., but there are far more with agronomic uses that are less well known and not yet developed commercially. Recent advances in technology and genomic resources provide tools to understand and realize the tremendous potential for using CAM crops to produce climate-resilient agricultural commodities in the future.

Elevated CO2 and temperature increase soil C losses from a soybean-maize ecosystem.

Warming temperatures and increasing CO2 are likely to have large effects on the amount of carbon stored in soil, but predictions of these effects are poorly constrained. We elevated temperature (canopy: +2.8 °C; soil growing season: +1.8 °C; soil fallow: +2.3 °C) for 3 years within the 9th-11th years of an elevated CO2 (+200 ppm) experiment on a maize-soybean agroecosystem, measured respiration by roots and soil microbes, and then used a process-based ecosystem model (DayCent) to simulate the decadal effects of warming and CO2 enrichment on soil C. Both heating and elevated CO2 increased respiration from soil microbes by ~20%, but heating reduced respiration from roots and rhizosphere by ~25%. The effects were additive, with no heat × CO2 interactions. Particulate organic matter and total soil C declined over time in all treatments and were lower in elevated CO2 plots than in ambient plots, but did not differ between heat treatments. We speculate that these declines indicate a priming effect, with increased C inputs under elevated CO2 fueling a loss of old soil carbon. Model simulations of heated plots agreed with our observations and predicted loss of ~15% of soil organic C after 100 years of heating, but simulations of elevated CO2 failed to predict the observed C losses and instead predicted a ~4% gain in soil organic C under any heating conditions. Despite model uncertainty, our empirical results suggest that combined, elevated CO2 and temperature will lead to long-term declines in the amount of carbon stored in agricultural soils.

Toward systems-level analysis of agricultural production from crassulacean acid metabolism (CAM): scaling from cell to commercial production.

Systems-level analyses have become prominent tools for assessing the yield, viability, economic consequences and environmental impacts of agricultural production. Such analyses are well-developed for many commodity crops that are used for food and biofuel, but have not been developed for agricultural production systems based on drought-tolerant plants that use crassulacean acid metabolism (CAM). We review the components of systems-level evaluations, and identify the information available for completing such analyses for CAM cropping systems. Specific needs for developing systems-level evaluations of CAM agricultural production include: improvement of physiological models; assessment of product processing after leaving the farm gate; and application of newly available genetic tools to the optimization of CAM species for commercial production.

A roadmap for research on crassulacean acid metabolism (CAM) to enhance sustainable food and bioenergy production in a hotter, drier world.

Crassulacean acid metabolism (CAM) is a specialized mode of photosynthesis that features nocturnal CO2 uptake, facilitates increased water-use efficiency (WUE), and enables CAM plants to inhabit water-limited environments such as semi-arid deserts or seasonally dry forests. Human population growth and global climate change now present challenges for agricultural production systems to increase food, feed, forage, fiber, and fuel production. One approach to meet these challenges is to increase reliance on CAM crops, such as Agave and Opuntia, for biomass production on semi-arid, abandoned, marginal, or degraded agricultural lands. Major research efforts are now underway to assess the productivity of CAM crop species and to harness the WUE of CAM by engineering this pathway into existing food, feed, and bioenergy crops. An improved understanding of CAM has potential for high returns on research investment. To exploit the potential of CAM crops and CAM bioengineering, it will be necessary to elucidate the evolution, genomic features, and regulatory mechanisms of CAM. Field trials and predictive models will be required to assess the productivity of CAM crops, while new synthetic biology approaches need to be developed for CAM engineering. Infrastructure will be needed for CAM model systems, field trials, mutant collections, and data management.

Development and use of bioenergy feedstocks for semi-arid and arid lands.

Global climate change is predicted to increase heat, drought, and soil-drying conditions, and thereby increase crop sensitivity to water vapour pressure deficit, resulting in productivity losses. Increasing competition between agricultural freshwater use and municipal or industrial uses suggest that crops with greater heat and drought durability and greater water-use efficiency will be crucial for sustainable biomass production systems in the future. Agave (Agavaceae) and Opuntia (Cactaceae) represent highly water-use efficient bioenergy crops that could diversify bioenergy feedstock supply yet preserve or expand feedstock production into semi-arid, abandoned, or degraded agricultural lands, and reclaim drylands. Agave and Opuntia are crassulacean acid metabolism species that can achieve high water-use efficiencies and grow in water-limited areas with insufficient precipitation to support traditional C3 or C4 bioenergy crops. Both Agave and Opuntia have the potential to produce above-ground biomass rivalling that of C3 and C4 crops under optimal growing conditions. The low lignin and high amorphous cellulose contents of Agave and Opuntia lignocellulosic biomass will be less recalcitrant to deconstruction than traditional feedstocks, as confirmed by pretreatments that improve saccharification of Agave. Refined environmental productivity indices and geographical information systems modelling have provided estimates of Agave and Opuntia biomass productivity and terrestrial sequestration of atmospheric CO2; however, the accuracy of such modelling efforts can be improved through the expansion of field trials in diverse geographical settings. Lastly, life cycle analysis indicates that Agave would have productivity, life cycle energy, and greenhouse gas balances comparable or superior to those of traditional bioenergy feedstocks, but would be far more water-use efficient.

Light to liquid fuel: theoretical and realized energy conversion efficiency of plants using crassulacean acid metabolism (CAM) in arid conditions.

There has been little attention paid to crassulacean acid metabolism (CAM) as a mechanism for bioenergy crop tolerance to water limitation, in part, because potential yields of CAM plants have been assumed to be lower than those of most commonly studied bioenergy crops. The photochemical efficiency, water-use efficiency (WUE), biomass production, and fuel yield potentials of CAM, C3, and C4 plants that are considered or already in use for bioenergy are reviewed here. The theoretical photosynthetic efficiency of CAM plants can be similar to or greater than other photosynthetic pathways. In arid conditions, the greater WUE of CAM species results in theoretical biomass yield potentials that are 147% greater than C4 species. The realized yields of CAM plants are similar to the theoretical yields that account for water-limiting conditions. CAM plants can potentially be viable commercial bioenergy crops, but additional direct yield measurements from field trials of CAM species are still needed.

Impact of different bioenergy crops on N-cycling bacterial and archaeal communities in soil.

Biomass production for bioenergy may change soil microbes and influence ecosystem properties. To explore the impact of different bioenergy cropping systems on soil microorganisms, the compositions and quantities of soil microbial communities (16S rRNA gene) and N-cycling functional groups (nifH, bacterial amoA, archaeal amoA and nosZ genes) were assessed under maize, switchgrass and Miscanthus x giganteus at seven sites representing a climate gradient (precipitation and temperature) in Illinois, USA. Overall, the site-to-site variation in community composition surpassed the variation due to plant type, and microbial communities under each crop did not converge on a 'typical' species assemblage. Fewer than 5% of archaeal amoA, bacterial amoA, nifH and nosZ OTUs were significantly different among these crops, but the largest differences observed at each site were found between maize and the two perennial grasses. Quantitative PCR revealed that the abundance of the nifH gene was significantly higher in the perennial grasses than in maize, and we also found significantly higher total N in the perennial grass soils than in maize. Thus, we conclude that cultivation of these perennial grasses, instead of maize, as bioenergy feedstocks can improve soil ecosystem nitrogen sustainability by increasing the population size of N-fixing bacteria.

Agave americana: Characteristics and Potential Breeding Priorities.

Agave americana L. is one of many Agave species that could be developed for the production of valuable agricultural products. Although all species in this genus use Crassulacean Acid Metabolism (CAM) and most have drought and heat tolerance, Agave americana also has the combined traits of high yield and cold tolerance. This review highlights key characteristics of Agave americana that make it an exceptional novel crop for fiber, sweeteners, bioproducts, and bioethanol with resilient traits for changing climate conditions. Then, it proposes potential directions for breeding that will support production in semi-arid climates. With selection and breeding, yields of 16 Mg ha-1 y-1 may be achieved. Current field observations, with no crop improvement, indicate ~9 Mg ha-1 is the maximum yield, and in arid regions, a yield of ~3 Mg ha-1 y-1 is observed. It may be beneficial to breed for a shorter time to flowering, as has been successful for Agave tequilana Weber var. azul, so that further breeding goals are achievable in a decadal timespan. Specific trait selection during breeding will depend on whether fiber or sugar yields are the desired products at a given location. Even without breeding, varieties of Agave americana are climate resilient alternatives for some current commodity crops.

Life-cycle analysis and the ecology of biofuels.

Biofuels have been proposed as an ecologically benign alternative to fossil fuels. There is, however, considerable uncertainty in the scientific literature about their ecological benefit. Here, we review studies that apply life-cycle analysis (LCA), a computational tool for assessing the efficiency and greenhouse gas (GHG) impact of energy systems, to biofuel feedstocks. Published values for energy efficiency and GHG differ significantly even for an individual species, and we identify three major sources of variation in these LCA results. By providing new information on biogeochemistry and plant physiology, ecologists and plant scientists can increase the accuracy of LCA for biofuel production systems.

Assessing hydrothermal carbonization as sustainable home sewage management for rural counties: A case study from Appalachian Ohio.

Improper management of home sewage treatment systems (HSTS) presents major challenges in the developing world, and even in many parts of developed countries, which contribute to health, environmental, economic, and social problems. Hydrothermal treatment, a thermochemical conversion process that is particularly useful for wet wastes, can produce fertilizers from septic tank wastes while eliminating human pathogens. However, hydrothermal treatment requires high temperature and high pressure, which might need additional economical justification when targeted for rural communities. The aim of this study was to investigate the economic feasibility of a hydrothermal treatment facility that can treat septic waste generated in Athens County, Ohio, a rural Appalachian county in the U.S., where failing HSTS have been observed quite frequently. Two different cases were considered for economic analysis, where Case I assumed a decentralized facility and Case II assumed a centralized facility. Results showed that both cases are economically feasible, where Case I and II reached breakeven in years 4 and 6, respectively. Additionally, despite the greater capital investment requirement for Case I, there was also a greater return on investment (ROI) of 2.85 compared to 1.52 for Case II. A sensitivity analysis was examined to determine the effect of solid content in either the septic tank or the reactor's feed, selling price of the fertilizer, and septage pumping costs on the project feasibility. The sensitivity analysis showed that pumping cost is the most significant factor affecting the project feasibility, while the ROI varied about ±80% and ± 200% for Case I and II, respectively, due to only about ±18% change in pumping cost.

Tissue Composition of Agave americana L. Yields Greater Carbohydrates From Enzymatic Hydrolysis Than Advanced Bioenergy Crops.

Agave americana L. is a highly productive, drought-tolerant species being investigated as a feedstock for biofuel production. Some Agave spp. yield crop biomass in semi-arid conditions that are comparable to C3 and C4 crops grown in areas with high rainfall. This study evaluates the bioethanol yield potential of A. americana by (1) examining the relationship between water use efficiency (WUE) and plant carbohydrates, (2) quantifying the carbohydrate and energy content of the plant tissue, and (3) comparing the products of enzymatic hydrolysis to that of other candidate feedstocks (Miscanthus x giganteus Greef et Deuter, Sorghum bicolor (L.) Moench, and Panicum virgatum L.). Results indicate that (1) WUE does not significantly affect soluble and insoluble (i.e., structural) carbohydrate composition per unit mass in A. americana; (2) without pretreatment, A. americana biomass had the lowest gross heat of combustion, or higher heating/calorific value, compared to high yielding C4 crops; and (3) after separation of soluble carbohydrates, A. americana cellulosic biomass was most easily hydrolyzed by enzymes with greater sugar yield per unit mass compared to the other biomass feedstocks. These results indicate that A. americana can produce substantial yields of soluble carbohydrates with minimal water inputs required for cultivation, and fiber portions of the crop can be readily deconstructed by cellulolytic enzymes for subsequent biochemical fermentation.

Predicting greenhouse gas emissions and soil carbon from changing pasture to an energy crop.

Bioenergy related land use change would likely alter biogeochemical cycles and global greenhouse gas budgets. Energy cane (Saccharum officinarum L.) is a sugarcane variety and an emerging biofuel feedstock for cellulosic bio-ethanol production. It has potential for high yields and can be grown on marginal land, which minimizes competition with grain and vegetable production. The DayCent biogeochemical model was parameterized to infer potential yields of energy cane and how changing land from grazed pasture to energy cane would affect greenhouse gas (CO2, CH4 and N2O) fluxes and soil C pools. The model was used to simulate energy cane production on two soil types in central Florida, nutrient poor Spodosols and organic Histosols. Energy cane was productive on both soil types (yielding 46-76 Mg dry mass · ha(-1)). Yields were maintained through three annual cropping cycles on Histosols but declined with each harvest on Spodosols. Overall, converting pasture to energy cane created a sink for GHGs on Spodosols and reduced the size of the GHG source on Histosols. This change was driven on both soil types by eliminating CH4 emissions from cattle and by the large increase in C uptake by greater biomass production in energy cane relative to pasture. However, the change from pasture to energy cane caused Histosols to lose 4493 g CO2 eq · m(-2) over 15 years of energy cane production. Cultivation of energy cane on former pasture on Spodosol soils in the southeast US has the potential for high biomass yield and the mitigation of GHG emissions.

How can land-use modelling tools inform bioenergy policies?

Targets for bioenergy have been set worldwide to mitigate climate change. Although feedstock sources are often ambiguous, pledges in European nations, the United States and Brazil amount to more than 100 Mtoe of biorenewable fuel production by 2020. As a consequence, the biofuel sector is developing rapidly, and it is increasingly important to distinguish bioenergy options that can address energy security and greenhouse gas mitigation from those that cannot. This paper evaluates how bioenergy production affects land-use change (LUC), and to what extent land-use modelling can inform sound decision-making. We identified local and global internalities and externalities of biofuel development scenarios, reviewed relevant data sources and modelling approaches, identified sources of controversy about indirect LUC (iLUC) and then suggested a framework for comprehensive assessments of bioenergy. Ultimately, plant biomass must be managed to produce energy in a way that is consistent with the management of food, feed, fibre, timber and environmental services. Bioenergy production provides opportunities for improved energy security, climate mitigation and rural development, but the environmental and social consequences depend on feedstock choices and geographical location. The most desirable solutions for bioenergy production will include policies that incentivize regionally integrated management of diverse resources with low inputs, high yields, co-products, multiple benefits and minimal risks of iLUC. Many integrated assessment models include energy resources, trade, technological development and regional environmental conditions, but do not account for biodiversity and lack detailed data on the location of degraded and underproductive lands that would be ideal for bioenergy production. Specific practices that would maximize the benefits of bioenergy production regionally need to be identified before a global analysis of bioenergy-related LUC can be accomplished.

Feedstocks for lignocellulosic biofuels.

In 2008, the world produced approximately 87 gigaliters of liquid biofuels, which is roughly equal to the volume of liquid fuel consumed by Germany that year. Essentially, all of this biofuel was produced from crops developed for food production, raising concerns about the net energy and greenhouse gas effects and potential competition between use of land for production of fuels, food, animal feed, fiber, and ecosystem services. The pending implementation of improved technologies to more effectively convert the nonedible parts of plants (lignocellulose) to liquid fuels opens diverse options to use biofuel feedstocks that reach beyond current crops and the land currently used for food and feed. However, there has been relatively little discussion of what types of plants may be useful as bioenergy crops.

Paediatricians and the pharmaceutical industry: an industry perspective of the challenges ahead.

The relationship between healthcare professionals and the pharmaceutical industry is under intense scrutiny. Accusations of corrupt practices have been levelled at the industry by both the professional and the lay press. The environment is changing, with rising expectations of transparency and ethical standards. In addition, society is becoming increasingly risk averse. There have been examples of poor practice by industry in the past, and industry is learning from these. Equally, there are many examples of excellent practice where industry has worked effectively and ethically with clinicians. The goal of industry is to bring new medicines to benefit patients and shareholders. Commercial success is dependent upon putting the patient at the centre of activity. It also allows re-investment into research and development. This allies industry and healthcare professionals with a common and key ethical arbitrator--the patient. Industry is changing as an acknowledgement of the need to adapt to the culture change demanded by society. Self regulation is increasingly active and transparent; guidelines, laws and internal/external regulators exist to examine industry. However, for the changes in regulations, behaviour and practices to be fully effective, there is a need for dialogue between industry and healthcare professionals. Commitment is needed from both sides to work together to manage the relationship.