Influence of Elevated CO2 and Temperature on the Photosynthesis and Respiration of White Clover Dependent on N2 Fixation

Single clonal plants of white clover (Trifolium repens L) grownfrom explants in a Perlite rooting medium, and dependent fornitrogen on N₂ fixation in root nodules, were grown for severalweeks in controlled environments which provided two regimesof CO₂, and temperature 23/18 °C day/night temperaturesat 680 µmol mol^–1 CO₂, (C680), and 20/15 °Cday/night temperatures at 340 µmol mol^–1 CO₂ (C340)After 3–4 weeks of growth, when the plants were acclimatedto the environmental regimes, leaf and whole-plant photosynthesisand respiration were measured using conventional infra-red gasanalysis techniques Elevated CO₂ and temperature increased ratesof photosynthesis of young, fully expanded leaves at the growthirradiance by 17–29%, despite decreased stomatal conductancesand transpiration rates Water use efficiency (mol CO₂ mol H₂O^–1)was also significantly increased Plants acclimated to elevatedCO₂, and temperature exhibited rates of leaf photosynthesisvery similar to those of C340 leaves ‘instantaneously’exposed to the C680 regime However, leaves developed in theC680 regime photosynthesised less rapidly than C340 leaves whenboth were exposed to a normal CO₂, and temperature environmentIn measurements where irradiance was varied, the enhancementof photosynthesis in elevated CO₂ at 23 °C increased graduallyfrom approx 10 % at 100 µmol m^–1 s^–1 to >27 % at 1170 µmol m^–2 s^–1 In parallel, wateruse efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 In parallel,water use efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 to approx100 % at the highest irradiance Elevated CO₂, and temperatureincreased whole-plant photosynthesis by > 40 %, when expressedin terms of shoot surface area or shoot weight No effects ofelevated CO₂ and temperature on rate of tissue respiration,either during growth or measurement, were established for singleleaves or for whole plants Dependence on N₂, fixation in rootnodules appeared to have no detrimental effect on photosyntheticperformance in elevated CO₂, and temperature Trifolium repens, white clover, photosynthesis, respiration, elevated CO₂, elevated temperature, water use efficiency, N₂ fixation     

The Strategy of Carbon Utilization in Uniculm Barley: II. THE EFFECT OF CONTINUOUS LIGHT AND CONTINUOUS DARK TREATMENTS

After a photoperiod of 8.25 h during which the youngest fullyexpanded leaf of uniculm barley plants was allowed to assimilate¹⁴CO₂ for 30 min, groups of plants were transfered either tocontinuous light or to continuous dark. Plants were harvestedover a 72 h period to examine the effect of the treatments (comparedwith control plants growing in normal light/dark cycles) onthe transport of ¹⁴C from the exposed leaf, the distributionof ¹⁴C assimilates to the rest of the plant, and the chemicalfate of assimilated ¹⁴C. In continuous light a substantial quantity (22% at 72 h) ofthe ¹⁴C assimilated by the leaf remained in that leaf in theform of starch and neutral sugars compared with only 4% in thecontrol fed leaf. Also the total amount of ¹⁴C respired fromplants maintained in continuous light was significantly less(c. 18% of the total originally fixed by 24 h) than that respiredfrom control plants (c. 36%). The result was that approximatelyequal amounts of ¹⁴C were accumulated in the growing leavesand roots of plants given continuous light or normal light/darkcycles. In continuous dark the fate of ¹⁴C was similar to that of controlplants. This is probably because the two treatments shared acommon light/dark environment for the first 22 h, during whichtime almost complete distribution and utilization of ¹⁴C occurred.     

The Respiratory Costs of Nitrogen Fixation in Soyabean, Cowpea, and White Clover: II. COMPARISONS OF THE COST OF NITROGEN FIXATION AND THE UTILIZATION OF COMBINED NITROGEN

Plants of soyabean, cowpea, and white clover were grown singlyin pots in Saxcil growth cabinets at 23/18 °C, 30/24 °C,and 20/15 °C, respectively, until seed maturation or for85 d (white clover). Two populations were produced within eachspecies: one population effectively nodulated and wholly dependentfor nitrogen on fixation in the root nodules, and a second populationcompletely lacking nodules but receiving abundant nitrate nitrogen.In each species, the two populations were compared in termsof rate of gross photosynthesis, rate of shoot respiration,and rate of root respiration. Source of nitrogen had littleor no effect on rate of photosynthesis or shoot respiration.In contrast, the rate of respiration of the nodulated rootsof plants fixing their own nitrogen was greater, sometimes two-foldgreater, than that of equivalent plants lacking nodules andutilizing nitrate nitrogen. This superiority in terms of rateof root respiration was generally confined to the period ofintense nitrogen fixation. An analysis of the magnitude of thisrespiratory burden in terms of daily photosynthesis indicatesthat, in all three legumes, plants fixing their own nitrogenrespire 11–13% more of their fixed carbon each day thanequivalent plants lacking nodules and utilizing nitrate nitrogen.     

Distribution patterns of assimilated 14C in vegetative and reproductive shoots of Lolium perenne and L. temulentum

The movement of ¹⁴C-labelled assimilate to the terminal meristem, stem, mature leaves, tillers and roots was measured in Loliurn perenn and Lolium temulentum after exposure to ¹⁴C0₂ of the youngest fully-expanded leaf and, on fewer occasions, the oldest healthy leaf on the main shoot. During early vegetative growth, the terminal meristem, tillers and roots received most of the ¹⁴C exported from the youngest leaf. As the shoot aged, more ¹⁴C was exported to the terminal meristem and tillers and less to roots. When the stem became a sizeable sink for ¹⁴C at the six-leaf (L. temulentum) or eleven-leaf (L. perenne) stage, less ¹⁴C moved to tillers and much less to roots. The terminal meristem continued to receive ¹⁴ at a steady rate throughout late vegetative growth. The transition from vegetative to reproductive growth in both species was marked by an abrupt increase in the export of ¹⁴C to stem from the upper leaf, but there was little change in the proportion of ¹⁴C which moved to the developing leaves and incipient inflorescence at the terminal meristem. At the same time, less ¹⁴C moved to tillers and much less to roots. Immediately before ear emergence, the export of ¹⁴C from the upper leaf (flag leaf) to the stem declined and the proportion moving to the ear increased, reaching a maximum of 55–75% as the ear emerged. The relative patterns of export of upper and lower leaves showed that while some ¹⁴ moved from each leaf to all meristems, the proximity of actively growing meristems appeared to be the main factor which determined the destination of most exported ¹⁴C. The distribution of ¹⁴C from upper and lower leaves was most alike in young vegetative plants of L. perenne. At later stages of development of both species, the terminal meristem and stem received most 14¹⁴C from the upper leaf, while roots and tillers received mos 14¹⁴C from the oldest leaf at the base of the shoot.     

A Quantitative Analysis of the Uptake of Carbon and of the Supply of 14C-labelled Assimilates to Areas of Meristematic Growth in Lolium temulentum

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby leaves on the main shoot to terminal meristem, stem, tillers,and roots was conducted during parallel periods of reproductiveand vegetative development in Lolium temulentum. The initial rate of entry of carbon into the shoot varied withthe area and photosynthetic efficiency of the assimilating leaf.Subsequently, respiratory losses of carbon during translocationand incorporation of assimilate at the site of utilization alsovaried. The combined effect of these differences resulted inthe supply of recently assimilated carbon being twofold greaterin reproductive shoots than in vegetative shoots, while withinshoots the carbon supply of the youngest fully expanded leafranged from four-or five-fold greater than the oldest leaf inyoung shoots, to two-or three-fold greater in older shoots.In both reproductive and vegetative shoots, the two or threeyoungest leaves thus dominated the supply of carbon for meristematicgrowth. Meristematic tissue in expanding leaves and leaf primordia atthe terminal meristem of the vegetative shoot received 18–27per cent of the total shoot carbon. This meristem utilized aboutthe same proportion of shoot carbon when it developed into aninflorescence, indicating no major change in the level of meristematicactivity. The proportion of shoot carbon utilized in stem growthincreased as both reproductive and vegetative shoots aged; thisincreased meristematic activity in stem internodes was accompaniedby reduced export of carbon to roots, which received less than10 per cent of the shoot carbon when the experiments ended.The main shoot translocated 20–30 per cent of its recentlyassimilated carbon to developing and rooted tillers, which assinks for carbon were thus as important as the terminal meristemand stem. This outward flow of carbon continued relatively uncheckedwhen donor and receptor shoots developed inflorescences.     

Studies on the Physiology of Flowering of Timothy (Phleum pratense L.) II. Influence of Daylength and Temperature on Size of the inflorescence

Inflorescence length in timothy increases when the photoperiodis reduced from 24 to 14 hours of light; it is also increasedby a reduction in ambient temperature from 75° to 55°F. There is a linear relation between total floret number andear length. Both factors affect ear length by influencing therate of growth of the spike between spikelet initiation andear emergence; this implies an effect on either the number ofprimary spikelet initials or the number of florets producedby branching, or both. Experiments with Lolium temulentum, wheredaylength and temperature influenced initiation and ear developmentin a way similar to that observed in timothy, suggest that thesefactors affect the number of florets at each primary initial.The interrelations of internal and external factors and theirinfluence on inflorescence size in the grasses is discussed.     

The Export and Distribution of 14C-labelled Assimilates from each Leaf on the Shoot of Lolium temulentum during Reproductive and Vegetative Growth

In both reproductive and vegetative plants of Lolium temulentumL., the export of ¹⁴C-labelled assimilates from each healthyleaf on the main shoot to terminal meristem, stem, tillers,and roots was measured each time a new leaf was expanded, fora period of 5 to 6 weeks. Some labelled assimilates moved fromeach leaf on the main shoot to every meristem in the same shoot,as well as to the tops and roots of adjacent organically attachedtillers. The terminal meristem of the reproductive shoot, which includedthe developing inflorescence, received 70–80 per centof the carbon assimilated by the emerged portion of the growingleaf, 15–25 per cent of the carbon assimilated by thetwo youngest expanded leaves, and 5–10 per cent of thatfrom each of the older leaves. A similar pattern of carbon supplyto the terminal meristem was found in vegetative shoots, exceptthat older leaves on young vegetative shoots supplied even lessof their carbon to the terminal meristem. The general conclusionis that developing leaves at the tip of the shoot receive aboutthe same proportion of carbon from each leaf as does a developinginflorescence. Young expanded leaves provided most labelled assimilates forstem growth; during both reproductive and vegetative growth,expanded leaves increased their export of labelled carbon tostem, and exported less of their ¹⁴C to roots and sometimesto tillers. In these reproductive and vegetative shoots, grown in a constantexternal environment, the major changes in the pattern of distributionof labelled assimilates appeared to be the result of increasedmeristematic activity in stem internodes; the development ofan inflorescence had no obvious direct effect on the carboneconomy of shoots.     

The Use of a Model to Investigate the Influence of some Environmental Factors on the Growth of Perennial Ryegrass

A modelof thegrasscrop isconstructed, inahichthe simulated valuesof crop morphology and physiology adapt automatically to changesin the environment. The model predicts photosynthesis, partitioningof assimilates and the growth or the cropin terms of leaves,stems and rmts from environmental data and the initial weightsof the crop components following defoliation. The model is usedto investigate the separate effects of light, temperature, canopystructure and maintenance respiration on the growth and denlopmcnt of the crop. It demonstrates how the crop attempts to optimizeaboveground growth, and the complexity of the interaction betweenthe physiological and environmental factors controlling growth. Lolium perenne, perennial ryegrass, photosynthesis, partition of assimilates, mathematical model