Acclimation of rice to changing atmospheric carbon dioxide concentration

Abstract. The effects were studied of season-long (75 and 88d) exposure of rice (Oryza sativa L. cv. IR-30) to a range of atmospheric CO₂ concentrations in outdoor, computer-controlled, environment chambers under natural solar radiation. The CO₂ concentrations were maintained at 160, 250, 330, 500, 660 and 900μmol mol^-1 air. Photosynthesis increased with increasing growth CO₂ concentrations up to 500u.mol moP1, but levelled off at higher CO₂ values. Specific leaf area also increased significantly with increasing CO₂. Although leaf dry weight and leaf area index increased, the overall response was not statistically significant. Leaf nitrogen content dropped slightly with elevated CO₂, but the response was not statistically significant. The specific activity of ribulose bisphosphate carboxylase/oxygenase (rubisco) declined significantly over the CO₂ concentration range 160 to 900μmol mol^-1. When expressed on a leaf area basis, rubisco activity decreased by 66%. This was accompanied by a 32% decrease in the amount of rubisco protein as a fraction of the total soluble leaf protein, and by 60% on a leaf area basis. For leaves in the dark, the total rubisco activity (CO₂/Mg²⁺-activated) was reduced by more than 60%. This indicates that rice accumulated an inhibitor in the dark, probably 2-car-boxyarabinitol 1-phosphate (CA-1-P). However, the inhibitor did not seem to be involved in the acclimation response. The degree of carbamylation of the rubisco enzyme was unchanged by the CO2 growth regime, except at 900 [μmol mol^-1 where it was reduced by 24%. The acclimation of rice to different atmospheric CO₂ conditions involved the modulation of both the activity and amount of rubisco protein in the leaf.     

Acclimation of the summer annual species, Lolium temulentum, to CO2 enrichment

Lolium temulentum L. Ba 3081 was grown hydroponically in air (350 μmol mol⁻¹ CO₂) and elevated CO₂ (700 μmol mol⁻¹ CO₂) at two irradiances (150 and 500 μmol m⁻² s⁻¹) for 35 days at which point the plants were harvested. Elevated CO₂ did not modify relative growth rate or biomass at either irradiance. Foliar carbon-to-nitrogen ratios were decreased at elevated CO₂ and plants had a greater number of shorter tillers, particularly at the lower growth irradiance. Both light-limited and light-saturated rates of photosynthesis were stimulated. The amount of ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco) protein was increased at elevated CO₂, but maximum extractable Rubisco activities were not significantly increased. A pronounced decrease in the Rubisco activation state was found with CO₂ enrichment, particularly at the higher growth irradiance. Elevated-CO₂-induced changes in leaf carbohydrate composition were small in comparison to those caused by changes in irradiance. No CO₂-dependent effects on fructan biosynthesis were observed. Leaf respiration rates were increased by 68% in plants grown with CO₂ enrichment and low light. We conclude that high CO₂ will only result in increased biomass if total light input favourably increases the photosynthesis-to-respiration ratio. At low irradiances, biomass is more limited by increased rates of respiration than by CO₂-induced enhancement of photosynthesis. Received: 23 February 1999 / Accepted: 15 June 1999     

Acclimation to temperature of the response of photosynthesis to increased carbon dioxide concentration in Taraxacum officinale

The relative stimulation of photosynthesis by elevated carbon dioxide in C₃ species normally increases strongly with increasing temperature. This results from the kinetic characteristics of Rubisco, and has potentially important implications for responses of vegetation to increasing atmospheric carbon dioxide. It is often assumed that because Rubisco characteristics are conservative, all C₃ species have the same temperature dependence of the response of photosynthesis to elevated carbon dioxide. However, in this field study of Taraxacum officinale, there were no significant differences in the relative stimulation of photosynthesis by elevated carbon dioxide among days with temperatures ranging from 15 to 34 °C. Nevertheless, short-term measurements indicated a strong temperature dependence of the stimulation. This suggested that acclimation to temperature caused the lack of variation in the seasonal data. Experiments in controlled environments indicated that complete acclimation of the relative stimulation of photosynthesis by elevated carbon dioxide occurred for growth temperatures of 10 – 25 °C. The apparent specificity of Rubisco for carbon dioxide relative to oxygen at 15 °C, as assayed in vivo by measurements of the carbon dioxide concentration at which carboxylation equalled oxygenation, also varied with growth temperature. Changes in the apparent specificity of Rubisco accounted for the acclimation of the temperature dependence of the relative stimulation of photosynthesis by elevated carbon dioxide. It is premature to conclude that low temperatures will necessarily reduce the relative stimulation of photosynthesis caused by rising atmospheric carbon dioxide. This revised version was published online in June 2006 with corrections to the Cover Date.     

The Developmental Anatomy of Leaves in Lolium temulentum

Measurements of the epidermal cells in Lolium temulentum andassessments of cell number indicate that differences in bladewidth are due mostly to variations in cell number, whereas changesin blade and sheath length result mainly from differences incell length. A marked increase in length of the flag-leaf aheath,however, was related to an increase in cell number. Considerable changes in the relative proportions of leaf bladeand sheath were observed in the flag leaf, and the leaves immediatelypreceding it, associated with inflorescence initiation. As oneconsequence of this the area of the flag leaf, the largest onthe shoot, is virtually constant under different environmentalcondi-tions. It is suggested that this aspect of correlateddevelopment is related to the nutrition of the panicle, sincein annual grasses such as the cereals the flag leaf may be responsiblefor producing up to 30 per cent. of the starch in the grain.     

The genetics of meiotic chromosome pairing in Lolium temulentum x Lolium perenne tetraploids

Summary The degree of preferential pairing of homologous chromosomes was estimated in a series of tetraploid hybrids of Lolium temulentum x Lolium perenne by means of cytological and genetic analyses. The correlations between the frequency of bivalents at first metaphase of meiosis in the hybrid tetraploids and the degree of preferential pairing calculated from the segregation pattern of isozyme alleles in a test cross was extremely high. The results showed clearly that suppression of heterogenetic pairing in these Lolium tetraploids is achieved by a genetic system involving the A chromosomes as well as the B chromosome system which has been known for some time. Certain similarities with the genetic system controlling pairing in polyploid wheats are discussed.     

Photosynthetic acclimation to rising atmospheric carbon dioxide concentration

With rising level of CO2 in the atmosphere plants are expected to be exposed to higher concentration of CO2. Since, CO2 is a substrate limiting photosynthesis particularly in C3 plants in the present atmosphere, the impact of elevated CO2 would depend mainly on how photosynthesis acclimates or adjusts to the long term elevated level of CO2. Photosynthetic acclimation is a change in photosynthetic efficiency of leaves due to long term exposure to elevated CO2. This change in photosynthetic efficiency could be a biochemical adjustment that may improve the overall performance of a plant in a high CO2 environment or it could be due to metabolic compulsions as a result of physiological dysfunction. Acclimation has generally become synonymous with the word response, if long term exposure to elevated CO2 decreases the photosynthesis rate (Pn) at a given CO2 level, it is called negative acclimation, if it stimulates Pn at a given CO2 level, it is called positive acclimation. Photosynthetic acclimation is clearly revealed by comparing Pn of ambient and elevated CO2 grown plants at same level of CO2. Species level differences in acclimation to elevated CO2 have been reported. The physiological basis of differential photosynthetic acclimation to elevated CO2 is discussed in relation to the regulation of photosynthesis and photosynthetic carbon partitioning at cellular level.     

Synaptonemal complex formation in hybrids of Lolium temulentum x Lolium perenne (L.)

Comparisons were made between two kinds of tetraploids derived from the hybrid Lolium temulentum x L. perenne. One hybrid behaves like an autotetraploid with multivalents at first metaphase of meiosis in pollen mother cells. The other behaves like an allotetraploid, in which pairing at first metaphase is restricted to bivalents comprised of strictly homologous chromosomes. The diploidisation of the latter form is controlled by determinants located on both the normal, A chromosomes and on supernumary B chromosomes. Reconstruction of synaptonemal complexes and their elements, from serial sections through pollen mother cell nuclei examined under the electron microscope, reveals that at zygotene pairing in both forms results in multivalent formation involving non-homologous as well as homologous chromosomes. The mechanism responsible for the diploidisation is, therefore, not based on a restriction of pairing at early meiosis to homologous chromosomes but on a correction or transformation of the multivalent chromosome associations to bivalents subsequent to zygotene. The transformation is not completed until late pachytene. In the multivalent-forming tetraploid a maximum of four chromosomes are associated at first metaphase. Yet configurations of a higher valency are found at zygotene. There is, therefore, a partial transformation of multivalents even in this autotetraploid form which restricts configurations at metaphase I to homologous and homoeologous chromosomes only. In both hybrids some homologous bivalents are not the product of resolution of multivalents but result from two-by-two pairing from the beginning of zygotene.     

Effects of enhanced ultraviolet radiation and carbon dioxide concentration on the moss Hylocomium splendens

In a laboratory experiment interaction effects of UV-B and CO₂ on photosynthesis and growth of the moss Hylocomium splendens were studied. The plants were exposed to two CO₂ levels (350 ppm and 600 ppm) and three UV-B levels (no UV-B, ambient UV-B and that corresponding to 20% ozone depletion) for 5 months. The effects were recorded by measuring the photosynthetic response and growth of the plants. There was a statistically significant change in photosynthetic efficiency and maximum photosynthetic rates due to time and to enhanced CO₂ concentration, whereas there was no effect due to UV-B. There was a decreased growth due to both UV-B and CO₂ and an interaction effect on growth (in length). The UV-B dose corresponding to the ambient level had a larger reducing effect on growth than the highest UV-B dose. This was a counter-intuitive result and the following tentative interpretation was made: differences in the measured UV-A/UV-B/PAR ratios between the treatments could explain the result provided there was a non-linear response to UV over the range of irradiance levels used.     

Sagebrush and grasshopper responses to atmospheric carbon dioxide concentration

Summary Seed- and clonally-propagated plants of Big Sagebrush (Artemisia tridentata var.tridentata) were grown under atmospheric carbon dioxide regimes of 270, 350 and 650 μl l⁻¹ and fed toMelanoplus differentialis andM. sanguinipes grasshoppers. Total shrub biomass significantly increased as carbon dioxide levels increased, as did the weight and area of individual leaves. Plants grown from seed collected in a single population exhibited a 3–5 fold variation in the concentration of leaf volatile mono- and sesquiterpenes, guaianolide sesquiterpene lactones, coumarins and flavones within each CO₂ treatment. The concentration of leaf allelochemicals did not differ significantly among CO₂ treatments for these seed-propagated plants. Further, when genotypic variation was controlled by vegetative propagation, allelochemical concentrations also did not differ among carbon dioxide treatments. On the other hand, overall leaf nitrogen concentration declined significantly with elevated CO₂. Carbon accumulation was seen to dilute leaf nitrogen as the balance of leaf carbon versus nitrogen progressively increased as CO₂ growth concentration increased. Grasshopper feeding was highest on sagebrush leaves grown under 270 and 650 μl l⁻¹ CO₂, but varied widely within treatments. Leaf nitrogen concentration was an important positive factor in grasshopper relative growth but had no overall effect on consumption. Potential compensatory consumption by these generalist grasshoppers was apparently limited by the sagebrush allelochemicals. Insects with a greater ability to feed on chemically defended host plants under carbon dioxide enrichment may ultimately consume leaves with a lower nitrogen concentration but the same concentration of allelochemicals. Compensatory feeding may potentially increase the amount of dietary allelochemicals ingested for each unit of nitrogen consumed.     

Growth, photosynthesis and assimilate partitioning in Lolium temulentum exposed to chilling temperatures

Vegetative plants of Lolium temulentum L, grown at 20°C with an 8 h photoperiod were transferred to either 5 or 2°C (8 h photoperiod) at 4th leaf maturity. Measurement of dry weight gain indicated a marked decline in relative growth rate below 5°C, with growth being reduced as much between 5 and 2°C as between 20 and 5°C. This reduction was not associated with increased mortality and was reversible if plants were returned to 20°C. Tissue explants from cold-treated plants retained the ability to extend if returned to 20°C Rates of extension in explants were less temperature sensitive than the relative growth rates observed in intact plants. Measurements of photosynthetic capacity, and of the patterns of accumulation of reserve carbohydrate in plants exposed to different temperatures, suggested that the inhibition of growth caused by chilling is not caused by an inability of the plants to fix sufficient carbon.     

Generation of transgenic Lolium temulentum plants by Agrobacterium tumefaciens-mediated transformation

Lolium temulentum L. (Darnel ryegrass) has been proposed to be used as a model species for functional genomics studies in forage and turf grasses, because it is a self-fertile, diploid species with a short life cycle and is closely related to other grasses. Embryogenic calluses were induced from mature embryos of a double haploid line developed through anther culture. The calluses were broken up into small pieces and used for Agrobacterium tumefaciens-mediated transformation. A. tumefaciens strain EHA105 harboring pCAMBIA1301 and pCAMBIA1305.2 vectors were used to infect embryogenic callus pieces. Hygromycin was used as a selection agent in stable transformation experiments. Hygromycin resistant calluses were obtained after 4–6 weeks of selection and transgenic plants were produced in 10–13 weeks after Agrobacterium-mediated transformation. Fertile plants were readily obtained after transferring the transgenics to the greenhouse. Transgenic nature of the regenerated plants was demonstrated by Polymerase chain reaction (PCR), Southern hybridization analysis, and GUS staining. Progeny analysis showed Mendelian inheritance of the transgenes. The transformation system provides a valuable tool for functionality tests of candidate genes in forage and turf grasses.