Arnold J. Bloom

Plant Responses to Multiple Environmental FactorsPhysiological ecology provides tools for studying how interacting environmental resources control plant growth

M ost plants require a similar balance of resources-energy, water, and mineral nutrients-to maintain optimal growth. Natural environments, however, differ by at least two orders of magnitude in the availability of these resources. Light intensity varies 100fold from the canopy to the floor of a rainforest (Bj6rkman 1981); annual precipitation ranges 500-fold (105000 mm/yr) from deserts to tropical rainforests; and the amount of nitrogen available to plants varies from 0.09 g/m2 * yr in polar desert (Dowding et al. 1981) to 22.8 g/m2 * yr in a rich tropical rainforest (Vitousek 1984). Plants growing in these diverse environments maintain tissue concen-

Photorespiratory Metabolism: Genes, Mutants, Energetics, and Redox Signaling

Photorespiration is a high-flux pathway that operates alongside carbon assimilation in C(3) plants. Because most higher plant species photosynthesize using only the C(3) pathway, photorespiration has a major impact on cellular metabolism, particularly under high light, high temperatures, and CO(2) or water deficits. Although the functions of photorespiration remain controversial, it is widely accepted that this pathway influences a wide range of processes from bioenergetics, photosystem II function, and carbon metabolism to nitrogen assimilation and respiration. Crucially, the photorespiratory pathway is a major source of H(2)O(2) in photosynthetic cells. Through H(2)O(2) production and pyridine nucleotide interactions, photorespiration makes a key contribution to cellular redox homeostasis. In so doing, it influences multiple signaling pathways, particularly those that govern plant hormonal responses controlling growth, environmental and defense responses, and programmed cell death. The potential influence of photorespiration on cell physiology and fate is thus complex and wide ranging. The genes, pathways, and signaling functions of photorespiration are considered here in the context of whole plant biology, with reference to future challenges and human interventions to diminish photorespiratory flux.

Increasing CO2 threatens human nutrition

Dietary deficiencies of zinc and iron are a substantial global public health problem. An estimated two billion people suffer these deficiencies, causing a loss of 63 million life-years annually. Most of these people depend on C3 grains and legumes as their primary dietary source of zinc and iron. Here we report that C3 grains and legumes have lower concentrations of zinc and iron when grown under field conditions at the elevated atmospheric CO2 concentration predicted for the middle of this century. C3 crops other than legumes also have lower concentrations of protein, whereas C4 crops seem to be less affected. Differences between cultivars of a single crop suggest that breeding for decreased sensitivity to atmospheric CO2 concentration could partly address these new challenges to global health.

Carbon dioxide enrichment inhibits nitrate assimilation in wheat and Arabidopsis.

Nitrate for Me, Ammonium for You The interdependence of plant nitrogen uptake and plant responses to carbon dioxide is well established, but the influence of inorganic nitrogen form—i.e., whether nitrate or ammonium—has been largely ignored. Bloom et al. (p. 899) present evidence from five independent methods in both a monocot and dicot species that carbon dioxide inhibition of nitrate assimilation is a major determinant of plant responses to rising atmospheric concentrations of carbon dioxide. This finding explains several phenomena, including carbon dioxide acclimation and decline in food quality. The large variation in these phenomena among species, locations, or years derives from the large variation in the relative dependence of plants on nitrate and ammonium as nitrogen sources among species, locations, or years. The relative importance of ammonium and nitrate for plant N nutrition in future cropping systems will be critical for quantity and quality of food. The inhibition of photorespiration by elevated carbon dioxide is paralleled by a reduction in the assimilation of nitrate. The concentration of carbon dioxide in Earth’s atmosphere may double by the end of the 21st century. The response of higher plants to a carbon dioxide doubling often includes a decline in their nitrogen status, but the reasons for this decline have been uncertain. We used five independent methods with wheat and Arabidopsis to show that atmospheric carbon dioxide enrichment inhibited the assimilation of nitrate into organic nitrogen compounds. This inhibition may be largely responsible for carbon dioxide acclimation, the decrease in photosynthesis and growth of plants conducting C3 carbon fixation after long exposures (days to years) to carbon dioxide enrichment. These results suggest that the relative availability of soil ammonium and nitrate to most plants will become increasingly important in determining their productivity as well as their quality as food.

Nitrogen assimilation and growth of wheat under elevated carbon dioxide.

Simultaneous measurements of CO2 and O2 fluxes from wheat (Triticum aestivum) shoots indicated that short-term exposures to elevated CO2 concentrations diverted photosynthetic reductant from NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} or NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{2}^{-}}}\end{equation*}\end{document} reduction to CO2 fixation. With longer exposures to elevated CO2, wheat leaves showed a diminished capacity for NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} photoassimilation at any CO2 concentration. Moreover, high bicarbonate levels impeded NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{2}^{-}}}\end{equation*}\end{document} translocation into chloroplasts isolated from wheat or pea leaves. These results support the hypothesis that elevated CO2 inhibits NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} photoassimilation. Accordingly, when wheat plants received NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} rather than NH\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{4}^{+}}}\end{equation*}\end{document} as a nitrogen source, CO2 enhancement of shoot growth halved and CO2 inhibition of shoot protein doubled. This result will likely have major implications for the ability of wheat to use NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} as a nitrogen source under elevated CO2.

Root Development and Absorption of Ammonium and Nitrate from the Rhizosphere

Plant roots operate in an environment that is extremely heterogeneous, both spatially and temporally. Nonetheless, under conditions of limited diffusion and against intense competition from soil microorganisms, plant roots locate and acquire vital nitrogen resources. Several factors influence the mechanisms by which roots respond to ammonium and nitrate. Nitrogen that is required for cell division and expansion derives primarily from the apex itself absorbing rhizosphere ammonium and nitrate. Root density and extension are greater in nutrient solutions containing ammonium than in those containing nitrate as the sole nitrogen source. Root nitrogen acquisition alters rhizosphere pH and redox potential, which in turn regulate root cell proliferation and mechanical properties. The net result is that roots proliferate in soil zones rich in nitrogen. Moreover, plants develop thinner and longer roots when ammonium is the primary nitrogen source, an appropriate strategy for a relatively immobile nitrogen form.

Wheat leaves emit nitrous oxide during nitrate assimilation

Nitrous oxide (N2O) is a key atmospheric greenhouse gas that contributes to global climatic change through radiative warming and depletion of stratospheric ozone. In this report, N2O flux was monitored simultaneously with photosynthetic CO2 and O2 exchanges from intact canopies of 12 wheat seedlings. The rates of N2O-N emitted ranged from <2 pmol⋅m−2⋅s−1 when NH\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{4}^{+}}}\end{equation*}\end{document} was the N source, to 25.6 ± 1.7 pmol⋅m−2⋅s−1 (mean ± SE, n = 13) when the N source was shifted to NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document}. Such fluxes are among the smallest reported for any trace gas emitted by a higher plant. Leaf N2O emissions were correlated with leaf nitrate assimilation activity, as measured by using the assimilation quotient, the ratio of CO2 assimilated to O2 evolved. 15N isotopic signatures on N2O emitted from leaves supported direct N2O production by plant NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} assimilation and not N2O produced by microorganisms on root surfaces and emitted in the transpiration stream. In vitro production of N2O by both intact chloroplasts and nitrite reductase, but not by nitrate reductase, indicated that N2O produced by leaves occurred during photoassimilation of NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{2}^{-}}}\end{equation*}\end{document} in the chloroplast. Given the large quantities of NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{3}^{-}}}\end{equation*}\end{document} assimilated by plants in the terrestrial biosphere, these observations suggest that formation of N2O during NO\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}{\mathrm{_{2}^{-}}}\end{equation*}\end{document} photoassimilation could be an important global biogenic N2O source.

Phosphate absorption: adaptation of tundra graminoids to a low temperature, low phosphorus environment

Seasonal changes in phosphate absorption rate were examined in three tundra graminoids growing under natural field conditions and in experimentally heated soils in the field at Barrow, Alaska. Although tundra plants exhibited a high optimum temperature for phosphate absorption similar to that of temperate plants, rates were relatively insensitive to short-term temperature change, suggesting that tundra plants routinely absorb phosphate from cold soil and do not depend upon daily or seasonal increases in soil temperature for phosphate acquisition. Variation in the calculated seasonal course of phosphate absorption resulted more from change in phosphate availability and phosphate status than from temperature acclimation. Absorption capacity was highest early in the growing season when root growth began and then decreased to a level which was maintained well into September, long after aboveground plant parts had senesced. Nearly half of the phosphate acquired by the plants investigated was probably acquired after shoots had begun a net translocation of nutrients belowground for winter storage. Plants acclimatized to a warm, anaerobic, nitrogenrich soil had a higher capacity for phosphate absorption but a lower affinity for phosphate than did control plants growing in cold soil. The applicability of laboratory-derived concepts of nutrient absorption to the seasonal dynamics of these processes in the field is discussed.

Root distribution in relation to soil nitrogen availability in field-grown tomatoes.

Tomato root growth and distribution were related to inorganic nitrogen (N) availability and turnover to determine 1) if roots were located in soil zones where N supply was highest, and 2) whether roots effectively depleted soil N so that losses of inorganic N were minimized. Tomatoes were direct-seeded in an unfertilized field in Central California. A trench profile/monolith sampling method was used. Concentrations of nitrate (NO3-) exceeded those of ammonium (NH4+) several fold, and differences were greater at the soil surface (0–15 cm) than at lower depths (45–60 cm or 90–120 cm). Ammonium and NO3- levels peaked in April before planting, as did mineralizable N and nitrification potential. Soon afterwards, NO3- concentrations decreased, especially in the lower part of the profile, most likely as a result of leaching after application of irrigation water. Nitrogen pool sizes and rates of microbial processes declined gradually through the summer.Tomato plants utilized only a small percentage of the inorganic N available in the large volume of soil explored by their deep root systems; maximum daily uptake was approximately 3% of the soil pool. Root distribution, except for the zone around the taproot, was uniformly sparse (ca. 0.15 mg dry wt g-1 soil or 0.5 cm g-1 soil) throughout the soil profile regardless of depth, distance from the plant stem, or distance from the irrigation furrow. It bore no relation to N availability. Poor root development, especially in the N-rich top layer of soil, could explain low fertilizer N use by tomatoes.

CO2 enrichment inhibits shoot nitrate assimilation in C3 but not C4 plants and slows growth under nitrate in C3 plants

The CO2 concentration in Earths atmosphere may double during this century. Plant responses to such an increase depend strongly on their nitrogen status, but the reasons have been uncertain. Here, we assessed shoot nitrate assimilation into amino acids via the shift in shoot CO2 and O2 fluxes when plants received nitrate instead of ammonium as a nitrogen source (deltaAQ). Shoot nitrate assimilation became negligible with increasing CO2 in a taxonomically diverse group of eight C3 plant species, was relatively insensitive to CO2 in three C4 species, and showed an intermediate sensitivity in two C3-C4 intermediate species. We then examined the influence of CO2 level and ammonium vs. nitrate nutrition on growth, assessed in terms of changes in fresh mass, of several C3 species and a Crassulacean acid metabolism (CAM) species. Elevated CO2 (720 micromol CO2/mol of all gases present) stimulated growth or had no effect in the five C3 species tested when they received ammonium as a nitrogen source but inhibited growth or had no effect if they received nitrate. Under nitrate, two C3 species grew faster at sub-ambient (approximately 310 micromol/mol) than elevated CO2. A CAM species grew faster at ambient than elevated or sub-ambient CO2 under either ammonium or nitrate nutrition. This study establishes that CO2 enrichment inhibits shoot nitrate assimilation in a wide variety of C3 plants and that this phenomenon can have a profound effect on their growth. This indicates that shoot nitrate assimilation provides an important contribution to the nitrate assimilation of an entire C3 plant. Thus, rising CO2 and its effects on shoot nitrate assimilation may influence the distribution of C3 plant species.