Hormoz BassiriRad

Root system adjustments: regulation of plant nutrient uptake and growth responses to elevated CO2

Nutrients such as nitrogen (N) and phosphorus (P) often limit plant growth rate and production in natural and agricultural ecosystems. Limited availability of these nutrients is also a major factor influencing long-term plant and ecosystem responses to rising atmospheric CO2 levels, i.e., the commonly observed short-term increase in plant biomass may not be sustained over the long-term. Therefore, it is critical to obtain a mechanistic understanding of whether elevated CO2 can elicit compensatory adjustments such that acquisition capacity for minerals increases in concert with carbon (C) uptake. Compensatory adjustments such as increases in (a) root mycorrhizal infection, (b) root-to-shoot ratio and changes in root morphology and architecture, (c) root nutrient absorption capacity, and (d) nutrient-use efficiency can enable plants to meet an increased nutrient demand under high CO2. Here we examine the literature to assess the extent to which these mechanisms have been shown to respond to high CO2. The literature survey reveals no consistent pattern either in direction or magnitude of responses of these mechanisms to high CO2. This apparent lack of a pattern may represent variations in experimental protocol and/or interspecific differences. We found that in addressing nutrient uptake responses to high CO2 most investigators have examined these mechanisms in isolation. Because such mechanisms can potentially counterbalance one another, a more reliable prediction of elevated CO2 responses requires experimental designs that integrate all mechanisms simultaneously. Finally, we present a functional balance (FB) model as an example of how root system adjustments and nitrogen-use efficiency can be integrated to assess growth responses to high CO2. The FB model suggests that the mechanisms of increased N uptake highlighted here have different weights in determining overall plant responses to high CO2. For example, while changes in root-to-shoot biomass allocation, r, have a small effect on growth, adjustments in uptake rate per unit root mass,

Changes in root NH4+ and NO3− absorption rates of loblolly and ponderosa pine in response to CO2 enrichment

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Biological Extreme Events: A Research Framework

, and photosynthetic N use efficiency, p*, have a significantly greater leverage on growth responses to elevated CO2 except when relative growth rate (RGR) reaches its developmental limit, maximum RGR (RGRmax).

Consequences of atmospheric nitrogen deposition in terrestrial ecosystems: old questions, new perspectives.

Centaurea maculosa, an invasive North American plant species, shows a high degree of tolerance to the root-boring biocontrol herbivore, Agapeta zoegana. For example, infested individuals of C. maculosa often exhibit more rigorous growth and reproduction compared with their non-infested counterparts. Compensatory responses to aboveground herbivores often involve increases in leaf area and/or photosynthetic capacity, but considerably less is known about root system compensatory responses to belowground herbivory. We used a 15N labeling approach to evaluate whether compensatory adjustments in N acquisition via changes in root morphology and/or physiological uptake capacity could explain the ability of C. maculosa to tolerate root herbivory. Root herbivory reduced whole plant N uptake by more than 30% and root uptake capacity by about 50%. Despite a marked reduction in N procurement, herbivory did not affect total biomass or shoot N status. Infested plants maintained shoot N status by shifting more of the acquired N from the root to the shoot. To our knowledge, shifting N allocation away from a root herbivore has not been reported and provides a plausible mechanism for the host plant to overcome an otherwise devastating effect of a root herbivore-induced N deficit.

Differential responses of tallgrass prairie species to nitrogen loading and varying ratios of NO3- to NH4+

Root growth and physiological uptake capacity for NH4+ and NO3− were examined for seedlings of loblolly and ponderosa pine grown for 160 days under two CO2 levels, ambient (35 Pa) and ambient plus 35 Pa (70 Pa). Fraction of biomass allocated to active fine roots as well as total N (NH4+ + NO3−) absorption per unit root dry mass were unaffected by CO2. On a whole-plant basis, elevated CO2 led to a significant increase in N acquisition in loblolly but not in ponderosa pine. However, even in loblolly pine where CO2 significantly increased plant N acquisition, the relative increase, in biomass far exceeded the gain in N, i.e. a 60% increase in total dry weight was accompanied by only a 30% increase in N gain in response to high CO2. We suggest that the commonly reported decline in tissue N concentration of these and other species at high CO2 is largely caused by inability of the root systems to sufficiently compensate for increased N demand. Elevated CO2 significantly altered root uptake capacity of the different N forms, i.e., high CO2 significantly increased NO3− absorption rates, but decreased NH4+ absorption rates in both species though the decrease in loblolly was insignificant. However, elevated CO2 increased root respiration rate in loblolly pine while significantly decreasing it in ponderosa pine. This indicates that CO2-induced changes in plant preference for inorganic N forms is not simply regulated by root energy status. If changes in plant preference for inorganic N forms represent typical responses to elevated CO2, the results could have important implications for N dynamics in managed and natural plant communities.

Effects of high atmospheric CO2 concentration on root hydraulic conductivity of conifers depend on species identity and inorganic nitrogen source

Models describing plant and ecosystem N cycles require an accurate assessment of root physiological uptake capacity for NH4+ and NO3- under field conditions. Traditionally, rates of ion uptake in field-grown plants are determined by using excised root segments incubated for a short period in an assay solution containing N either as a radioactive or stable isotope tracer (e.g., 36ClO3 as a NH4+ analogue, 14CH3NH3 as an NO3- analogue or 15NH4+ and 15NO3-). Although reliable, this method has several drawbacks. For example, in addition to radioactive safety issues, purchase and analysis of radioactive and stable isotopes is relatively expensive and can be a major limitation. More importantly, because excision effectively interrupts exchange of compounds between root and shoot (e.g., carbohydrate supply to root and N transport to shoot), the assay must be conducted quickly to avoid such complications. Here we present a novel field method for simultaneous measurements of NH4+ and NO3- uptake kinetics in intact root systems. The application of this method is demonstrated using two tree species; red maple (Acer rubrum) and sugar maple (Acer saccharum) and two crop species soybean (Glycine max) and sorghum (Sorghum bicolor). Plants were grown in open-top chambers at either ambient or elevated levels of atmospheric CO2 at two separate US national sites involved in CO2 research. Absolute values of net uptake rates and the kinetic parameters determined by our method were found to be in agreement with the literature reports. Roots of the crop species exhibited a greater uptake capacity for both N forms relative to tree species. Elevated CO2 did not significantly affect kinetics of N uptake in species tested except in red maple where it increased root uptake capacity, V, for NH4+. The application, reliability, advantages and disadvantages of the method are discussed in detail.

Root System Characteristics and Control of Nitrogen Uptake

Efforts designed to understand and predict adaptation responses of organisms and populations to global climate change must make a clear distinction between responses to changes in average conditions (e.g., doubling of atmospheric carbon dioxide concentration accompanied by an average increase of 1°–3°C in global air temperature by the end of this century) and responses resulting from increased incidence of extreme events [Loehle and LeBlanc, 1996; Easterling et al., 2000; Garrett et al., 2006]. Such distinction is critical because, unlike changes in average conditions, extremes (e.g., megadroughts, fire, flooding, hurricanes, heat waves, and pest outbreaks) are typically short in duration but challenge organisms and populations considerably further beyond their ability to acclimate than those expected from average trends in climate changes.

Coordinated responses of plant hydraulic architecture with the reduction of stomatal conductance under elevated CO2 concentration

effects on ecosystem processes (Arens et al. 2008). It must also be noted that in regions where N deposition has stabi-lized or declined, the legacy effects of decades of N dep-osition are likely to override the declining rates of depo-sition, but little is known about such effects. Secondly, atmospheric N deposition in many regions of the world, including China and India (Vet et al. 2014; Liu et al. 2013), and most likely Brazil(Allen et al. 2011), is on the rise. These regions are home to some of the most productive and diverse ecosystems of the world whose responses to N dep-osition are still unknown. Third, even in regions where N deposition has been on the decline, the decrease is predom-inantly in the oxidized (NO