The Responses of Net Photosysthesis to Light and Temperature in Spartina townsendii(sensu lato), a C4 Species from a Cool Temperate Climate

Carbon dioxide and water vapour exchanges for single attachedleaves of the temperate C₄ grass Spartina townsendii were measuredunder controlled environment conditions in an open gas-exchangesystem. The responses of net photosynthesis, stomatal resistance,and residual resistance to leaf temperature and photon fluxdensity are described. The light and temperature responses ofnet photosynthesis in S. townsendii are compared to informationon these responses in both temperate C₃ grasses and sub-tropicalC₄ grasses. Adaptation of photosynthesis in this C₄ speciesto a cool temperate climate is indicated both by the light andtemperature responses of net photo-synthesis. Unlike the C₄grasses examined previously, significant rates of net photosynthesiscan be detected at leaf temperatures below 10?C. Rates of netphotosynthesis equal or exceed those reported for temperateC₃ grasses at all of the temperature (5–40?C) and photonflax density (13–2500µmol m^–2 s^–1) conditionsexamined. Maximum rates of net photosynthesis in S. townsendiiare almost double those reported for C₃ herbage grasses. Unliketemperate C₃ grasses, the major limitation to net photosynthesisat low leaf temperatures (10?C and below) is the stomatal resistance,showing that the low residual resistance characteristic of C₄species is maintained in S. townsendii even at low leaf temperatures.     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions II. Nitrogen Fixation and Nitrate Uptake

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and harvested at intervals over and88 d growht period. The swards received a nutrient solutiondaily, which was either High (220 mg l^–1) or Low (10 mgl^–1) in nitrate N. The nitrate was labelled with the ¹⁵Nisotope. An acetylene reduction assay was carried out on eachsward just prior to harvest. Rates of acetylene reduction agreed qualitatively with the ^l5Nanalyses but absolute values did not match (assuming a 4:1 C₂H₄:N₂ratio) and errors in the acetylene assay are discussed. In theLow-N swards clover relied almost entirely on symbioticallyfixed N₂, fixing more than ten times as much as the High-N cloverplants. In the Low-N treatment the grass was N-deficient despiteobtaining much more nitrate per unit root dry weight than clover.In the High-N swards, however, clover took up more nitrate perunit root weight than grass. The High-N clover plants also fixedsome N₂ and maintained a higher total-N content than grass throughoutthe period. There was no evidence of transfer of symbioticallyfixed N from the clover to the grass in either treatment. Trifolium repens, Lolium perenne, nitrate, nitrogen fixation, ¹⁵N, acetylene reduction     

The Seasonal Pattern of Growth and Production of a Temperate C4 Species, Cyperus longus

Growth and production of the temperate C₄ species Cyperus longusL. was measured throughout a growing season in an establishedplot in Eastern Ireland. The maximum standing live biomass reachedwas 2·5 kg m^–2. Estimates of unit leaf rate (ULR)and leaf area index (LAI) were made. The product of these quantitiesgave the crop growth rate (CGR) each week. C. longus was foundto maintain high values of LAI throughout the summer, with amaximum value of about 13 in early August. CGR reached a peakin early July. The optimum LAI was 11·6. Temperaturesat five levels in the plant canopy, and the amount of solarradiation intercepted by the canopy were measured continuouslyduring the summer. The mean daily rate of leaf extension waspositively correlated with the mean daily air temperature abovethe canopy but the temperature coefficient of the process waslow compared with other temperate species. The percentage ofsolar radiation intercepted by the canopy increased rapidlyin early summer, and canopy closure had occurred by mid-June.Rates of net photosynthesis were measured on young and old leafmaterial in situ at the time of peak LAI. In young leaves themaximum rates of net photosynthesis were higher than those publishedfor a range of temperate C₃ species, but similar to those foundin another temperate C₄ species, Spartina townsendii. Key words: C₄ photosynthesis, leaf growth, productivity     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions I. Carbon Assimilation and Utilization

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and their growth and carbon economyexamined. The swards received a nutrient solution daily, whichcontained either High (220 mg l^–1) or Low (10 mg l^–1)nitrate N. Rates of canopy photosynthesis and respiration, and final drymatter yields were similar in the two treatments although theproportions of grass and clover differed greatly. The Low-Nswards were made up largely of clover. The grass plants in theseswards had high root: shoot ratios and low relative photosyntheticrates – both signs of N deficiency – and were clearlyunable to compete with the vigorously growing Low-N clover plants.These had higher relative growth rates and dry matter yieldsthan their High-N counterparts. In the High-N swards clovercontributed around 50 per cent to the sward dry weight throughoutthe measurement period despite having a smaller proportion ofits dry weight in photosynthetic tissue (laminae) than grassover much of it. The latter was compensated for, initially bya higher specific leaf area than grass, and later by a higherphotosynthetic rate per unit leaf weight. The results are discussedin relation to observed declines in the clover content of swardsafter the addition of nitrogen fertilizer in the field. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, nitrogen, photosynthesis, carbon balance     

The Effects of Temperature on Leaf Growth in Cyperus longus, a Temperate C4 Species

Plants of the C₄ sedge Cyperus longus L. were grown at 10, 20and 30 °C. An asymptotic growth curve, the Richards function,was fitted to growth data for successive leaves. The mean rateof leaf appearance was a linear function of temperature with0.014 leaves appearing per day for every 1 °C increase intemperature. The instantaneous relative rate of leaf extensionshowed a marked ontogenetic drift which was most rapid at 30°C and slowest at 10 °C. The mean absolute extensionrate for foliage had a temperature coefficient of 0.16 cm d^–1° C^–1 in the range from 10 to 30 °C. The durationof leaf growth was independent of leaf number at 10 and 20 °Cbut increased linearly with leaf number at 30 °C. The smalldifferences in relative growth rate at the three temperaturesresulted in large differences in foliage area produced at theend of a 30 d growth period. The final foliage areas at 20 and10 °C were 51 and 9% respectively of that at 30 °C. Cyperus longus, temperature, leaf growth, Richards function, growth analysis     

Gas Exchange and Carbohydrate and Nitrogen Concentrations in Leaves of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) at Different Carbon Dioxide Concentrations and Temperatures

Pascopyrum smithii (C₃) andBouteloua gracilis (C₄) are importantforage grasses native to the Colorado shortgrass steppe. Thisstudy investigated photosynthetic responses of these grassesto long-term CO₂enrichment and temperature in relation to leafnonstructural carbohydrate (TNC) and [N]. Glasshouse-grown seedlingswere transferred to growth chambers and grown for 49 d at twoCO₂concentrations (380 and 750 µmol mol^-1) at 20 and 35°C, and two additional temperatures (25 and 30 °C) at750 µmol mol^-1CO₂. Leaf CO₂exchange rate (CER) was measuredat a plant's respective growth temperature and at two CO₂concentrationsof approx. 380 and 700 µmol mol^-1. Long-term CO₂enrichmentstimulated CER in both species, although the response was greaterin the C₃,P. smithii . Doubling the [CO₂] from 380 to 750 µmolmol^-1stimulated CER ofP. smithii slightly more in plants grownand measured at 30 °C compared to plants grown at 20, 25or 35 °C. CO₂-enriched plants sometimes exhibited lowerCER when compared to ambient-grown controls measured at thesame [CO₂], indicating photosynthetic acclimation to CO₂growthregime. InP. smithii , such reductions in CER were associatedwith increases in TNC and specific leaf mass, reductions inleaf [N] and, in one instance, a reduction in leaf conductancecompared to controls. InB. gracilis , photosynthetic acclimationwas observed more often, but significant changes in leaf metabolitelevels from growth at different [CO₂] were generally less evident.Temperatures considered optimal for growth (C₃: 20 °C; C₄:35 °C) sometimes led to CO₂-induced accumulations of TNCin both species, with starch accumulating in the leaves of bothspecies, and fructans accumulating only inP. smithii. Photosynthesisof both species is likely to be enhanced in future CO₂-enrichedand warmer environments, although responses will sometimes beattenuated by acclimation. Acclimation; blue grama (Bouteloua gracilis (H.B.K.) Lag ex Steud.); leaf nitrogen concentration; nonstructural carbohydrates; photosynthesis; western wheatgrass (Pascopyrum smithii (Rydb.) Love)     

The Potential of Two Perennial C4 Grasses and a Perennial C4 Sedge as Ligno-cellulosic Fuel Crops in N.W. Europe. Crop Establishment and Yields in E. England

Three perennial C₄ rhizomatous species, Cyperus longus L., Spartinacynosuroides and Spartina pectinata Link, were examined as potentialrenewable energy crops. These species are unusual among C₄ plantsin showing natural distributions which extend into cool temperateregions. This study examined whether these species could beestablished in the cool temperate climate of eastern Englandand whether they could consistently attain the relatively highdry matter yields associated with C₄ plants of warmer regions.Clonally produced material was planted in 1986, on two siteswith contrasting soil types in Essex, eastern England. Plantingwas within a randomized-block design incorporating replicatedplots of each species, both with and without fertilizer. Survivorshipand stem demography were monitored at monthly intervals from1986 to Jun. 1989 for stem recruitment and to Dec. 1991 forstem density. Yields were determined from 1987 (the year followingestablishment) to 1993. Survivorship of the planted propagules over the first 12 monthswas 92% for S. pectinata , 96% for S. cynosuroides and 100%for C. longus. Recruitment of new stems peaked in Apr. of mostyears, although a significant number of new stems appeared asearly as Feb. Stem death peaked in Sep. or early Oct. and allabove-ground stems had died by mid-Nov. Stem density trendsindicated that 2-4 years were required to reach a steady-statedensity, depending upon species. The stem density of the twoSpartina species had reached more than 1000 m^-2 in 1989 althoughthat of S. pectinata fluctuated considerably in the subsequentyears. C. longus stem density rose to approx. 600 m^-2 by 1988and did not change significantly in the subsequent years. In the 6 years following establishment, annual yields averagedacross all fertilizer treatments and both sites were 1·0,1·1 and 1·3 kg m^-2 for C. longus, S. cynosuroidesand S. pectinata, respectively. The average annual yield ofall three species at the site with the heavier soil was 1·3kg m^-2. This was significantly greater than the 1·0 kgm^-2 on the lighter soil. Nitrogen addition did not significantlyincrease yield. Even in the absence of any nitrogen addition,the annual yield of S. pectinata averaged 1·2 kg m^-2over the 6 years, with no evidence of any decline with the increasingage of the stands.Copyright 1995, 1999 Academic Press Cyperus longus, Spartina cynosuroides, Spartina pectinata, energy crop, dry matter yield     

Effect of Elevated CO2 on Stomatal Size and Distribution in Perennial Ryegrass

Plants of ryegrass (Lolium perenne L. cv. Melle) were grownfrom the early seedling stage in growth cabinets at a day/nighttemperature of 20/15 °C, with a 12-h photoperiod, and aCO₂ concentration of either 340 or 680 ± 15 µl1^–1 CO₂. Young, fully-expanded, acclimated leaves fromprimary branches were sampled for length of stomata, and ofepidermal cells between stomata, numbers of stomata and epidermalcells per unit length of stomatal row, numbers of stomatal rowsacross the leaf and numbers of stomatal rows between adjacentvein ridges. Elevated CO₂ had no significant effect on any ofthe measured parameters. Elevated CO₂, Lolium perenne, ryegrass, stomatal distribution, stomatal size     

Effect of Temperature and Nitrogen Supply on the Growth of Perennial Ryegrass and White Clover. 1. Carbon and Nitrogen Economies of Mixed Swards at Low Temperature

Simulated mixed swards of perennial ryegrass (Lolium perenneL. cv. S23) and white clover (Trifolium repens L. cv. S100)were grown from seed under a constant 10°C day/8°C nighttemperature regime and their growth, and carbon and nitrogeneconomies examined. The swards received a nutrient solution,every second day, which contained either high (220 µgg^–1) or low (40 µg g^–1) nitrate N. The High-N swards had rates of canopy photosynthesis and drymatter production (over the linear phase of growth) similarto those previously shown by mixed swards at high temperature.The Low-N swards grew more slowly; canopy photosynthesis, ata given LAI, was similar to that at High-N but lower LAI's weresustained. Clover increased its contribution to total carbonuptake and total dry weight throughout the period in the Low-Ntreatment and, despite the fact that grass took up most of theavailable nitrate, clover maintained a consistently higher Ncontent by virtue of N₂-fixation. At High-N, grass dominated throughout the measurement period.Earlier, when plants grew as spaced individuals, clover grewless well than grass, but once the canopy was closed it hada similar relative growth rate and thus maintained a steadyproportion of total sward dry weight. It is proposed that earlyin the development of the crop, leaf area production is thelimiting factor for growth, and that in this respect cloveris adversely affected by low temperature relative to grass.Later, as the LAI of the crop builds up, and the canopy becomesfully light intercepting, net canopy photosynthesis plays amore dominant role and here the higher photosynthetic rate perunit leaf area of the clover is crucial. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, low temperature, nitrogen, photosynthesis     

Response to Selection for Dark Respiration Rate of Mature Leaves in Lolium perenne and its Effects on Growth of Young Plants and Simulated Swards

Selections for slow and for fast rate of dark respiration ofmature leaves were made from within Lolium perenne cv. S23.Selected parents were pair-crossed to provide 15 F₁ familieswith slow respiration and 15 with fast. Dark respiration was inherited and families with contrastingrates were subjected to sequentially harvested growth analysesfrom the third leaf stage to that of 95 per cent light interceptionin a growth room. Seven periods of regrowth of simulated swardsof the families were then recorded. During development of theprimary canopy, growth of the selections did not differ untilthe final harvest interval. At this stage slow respiration familieshad faster (P < 0.05) net assimilation rate and greater plantdry weight (P < 0.05) and leaf area index (P < 0.05) thanthe fast respiration group. Relative growth rate followed thesame trend. In the swards after each regrowth dry matter yieldof the slow respiration group was greater than that of the fast. In another experiment, simulated swards of six slow respirationfamilies yielded more than swards of six fast respiration familiesover sequential regrowth periods in a glasshouse from May toNovember: S23 was intermediate. Differences were most duringAugust and September. Crop growth rate at each harvest correlated(P < 0.05 or P < 0.01) with previously determined leafrespiration at 25 °C. Leaf protein levels in August weaklycorrelated (r = +0.57, P < 0.05) with respiration rate perunit dry weight but there was a significant residual negativecorrelation (r = –0.67, P < 0.05) between the rateper unit protein and growth at that time. Results are discussedin relation to the concept of ‘maintenance-relàted’respiration. Lolium perenne L., perennial ryegrass, respiration, maintenance respiration, relative growth rate, leaf area ratio     

The Effects of Temperature and Form of Nitrogen Supply on the Relative Contribution of Root Uptake and Remobilization in Supplying Nitrogen for Laminae Regrowth of Lolium perenne L.

Plants of Lolium perenne L. cv. S23 were grown in sand culturesupplied with either ammonium (NH₄⁺) or nitrate (NO₃^–)in an otherwise complete nutrient solution at 12°C or 20°C.Three weeks after germination, plants were clipped weekly tosimulate grazing. After 10 weeks growth all nitrogen (N) wassupplied enriched with ¹⁵N to quantify the effects of form ofN supply and temperature on the relative ability of currentroot uptake and remobilization to supply N for laminae regrowth. The form of N supply had no effect on the dry matter partitioning,while at 20°C more dry weight was allocated to laminae regrowthand less to the remaining plant material. The current root uptakeof N, which subsequently appeared in the laminae regrowth, wassimilar for plants supplied with NH₄⁺ or NO₃^–, and bothwere equally reduced at the lower temperature of growth. Remobilizationof N to laminae regrowth was greater for plants receiving NH₄⁺than NO₃^–; remobilization with either form of N supplywas reduced at the lower temperature of growth. Remobilizationwas reduced to a lesser extent at 12°C than current rootuptake. It was concluded that remobilization became relativelymore important in supplying N for regrowth of laminae at lowertemperatures. Key words: Lolium perenne, ammonium, nitrate, temperature, remobilization     

Carbon Exchange Rate of Intact Individual Soya Bean Pods. 2. Ontogeny of Temperature Sensitivity

Diurnal temperature fluctuations induced change in soya bean-pod[Glycine max (L.) Merr.] carbon exchange rate (CER, where positiveCER represents CO₂ evolution). CER appeared to depend linearlyon temperature. Linear regressions of CER on temperature interceptedthe temperature axis at 5°C (i.e. zero CER at 5°C).Slopes of these regressions (i.e. temperature sensitivity) changedover the season. The CER-temperature sensitivity coefficient,K, (calculated from observed values of CER. pod temperatureand temperature intercept) rose from less than 0·02 mgCO₂ h^–1 pod^–1 °C^–1 during early pod-flll,peaked at over 0·04 mg CO₂ h^–1 pod^–1 °C^–1at mid pod-fill, and then declined during late pod-fill andmaturation. Glycine max (L.) Merr., Soya bean, carbon exchange rate, temperature     

Nitrate Effects on Growth of the First Four Main Stem Leaves of a Range of Temperate Cereals and Pasture Grasses

The effects of different applied NO₃^– concentrations onextension growth and final length and area of leaves 1–4of five cereals and six pasture grasses of temperate originwere examined. Increased applied NO₃^– in the range 0.1–0.5.0mol m^–3 caused decreased duration of growth but increasedgrowth rate and final length of leaves 2–4 of the cerealsAvena saliva, Hordeum vulgare, Secale cereale, x Triticosecaleand Triticum aestivum. For all cereals, increased NO₃^–resulted in increased area of leaves 1–4. Pasture grasseswere supplied either 0.5 or 50 mol m^–3 NO₃^–. Increasedapplied NO₃^– (0.5–5.0 mol m^–3) resulted indecreased duration of growth and increased growth rate and finalarea of leaves 1–4 of Bromus wiltdenowii, leaves 2–4ofFestuca arundinaceae and leaves 3 and 4 of Lolium muitiflorum.In addition, length of leaves 3 and 4 of B. witidenowii increasedwith increased NO₃^–. Increased NO₃^– resulted inincreased area of leaves 2–4 of Dactylis gtomerata andLolium perenne and leaves 3 and 4 of Phalaris aquaiica but hadno effect on extension growth of all three species. Avena sativa L, oat, Hordeum vulgare L, barley, Secale cereale L, rye, x Triticosecale Wittm, triticale, Triticum aestivum L, wheat, Bromus willdenowii Kunth, prairie grass, Dactylis gtomerata L, cocksfoot, Festuca arundinaceae Shreb, tall fescue, Lolium multijlorum Lam, Italian ryegrass, Lolium perenne L, perennial ryegrass, Phalaris aquatica L, nitrate, leaf extension, leaf expansion     

Growth and Partitioning in Pascopyrum smithii (C3) and Bouteloua gracilis(C4) as Influenced by Carbon Dioxide and Temperature

This study investigated how CO₂and temperature affect dry weight(d.wt) accumulation, total nonstructural carbohydrate (TNC)concentration, and partitioning of C and N among organs of twoimportant grasses of the shortgrass steppe,Pascopyrum smithiiRydb. (C₃) andBouteloua gracilis(H.B.K.) Lag. ex Steud. (C₄).Treatment combinations comprised two temperatures (20 and 35°C)at two concentrations of CO₂(380 and 750 µmol mol^-1),and two additional temperatures of 25 and 30°C at 750 µmolmol^-1CO₂. Plants were maintained under favourable nutrient andsoil moisture and harvested following 21, 35, and 49d of treatment.CO₂-induced growth enhancements were greatest at temperaturesconsidered favourable for growth of these grasses. Comparedto growth at 380 µmol mol^-1CO₂, final d.wt of CO₂-enrichedP.smithiiincreased 84% at 20°C, but only 4% at 35°C. Finald.wt ofB. graciliswas unaffected by CO₂at 20°C, but wasenhanced by 28% at 35°C. Root:shoot ratios remained relativelyconstant across CO₂levels, but increased inP. smithiiwith reductionin temperature. These partitioning results were adequately explainedby the theory of balanced root and shoot activity. Favourablegrowth temperatures led to CO₂-induced accumulations of TNCin leaves of both species, and in stems ofP. smithii, whichgenerally reflected responses of above-ground d.wt partitioningto CO₂. However, CO₂-induced decreases in plant tissue N concentrationswere more evident forP. smithii. Roots of CO₂-enrichedP. smithiihadgreater total N content at 20°C, an allocation of N below-groundthat may be an especially important adaptation for C₃plants.Tissue N contents ofB. graciliswere unaffected by CO₂. Resultssuggest CO₂enrichment may lead to reduced N requirements forgrowth in C₃plants and lower shoot N concentration, especiallyat favourable growth temperatures. Acclimation to CO₂; blue grama; Bouteloua gracilis ; carbohydrate; climate change; global change; grass; growth; growth temperature optima; nitrogen; N uptake; Pascopyrum smithii; western wheatgrass     

Nitrate Effects on Growth of the First Four Main Stem Leaves of a Range of Temperate Cereals and Pasture Grasses

The effects of different applied NO₃^– concentrations onextension growth and final length and area of leaves 1–4of five cereals and six pasture grasses of temperate originwere examined. Increased applied NO₃^– in the range 0.1–50mol m^–3; caused decreased duration of growth but increasedgrowth rate and final length of leaves 2–4 of the cerealsAvena saliva, Hordeum vulgare, Secale cereale x Triticosecaleand Triticum aestivum. For all cereals, increased NO₃^–resulted in increased area of leaves 1-4. Pasture grasses weresupplied either 0.5 or 50 mol m^–3; NO₃^–. Increasedapplied NO₃^– (0.5–50 mol m^–3) resulted indecreased duration of growth and increased growth rate and finalarea of leaves 1–4 of Bromus willdenowii leaves 2–4of Festuca arundinaceae and leaves 3 and 4 of Lolium multiflorum.In addition, length of leaves 3 and 4 of B. willdenowii increasedwith increased NO₃. Increased NO₃ resulted in increased areaof leaves 2–4 of Daciylis glomerata and Lolium perenneand leaves 3 and 4 of Phalaris aquatica but had no effect onextension growth of all three species. Avena saliva L., oat, Hordeum vulgare L., barley, Secale cereaie L., rye, x Triticosecale Wittm, triticale, Triticum aestivum L., wheat, Bromus willdenowii Kunth, prairie grass, Dactylis glomerata L., cocksfoot, Festuca arundinaceae Shreb, tall fescue, Lolium multiflorum Lam, Italian ryegrass, Lolium perenne L, perennial ryegrass, Phalaris aquatica L, nitrate,, leaf extension, leaf expansion     

Contrasting CO2 and temperature effects on leaf growth of perennial ryegrass in spring and summer

The effects of increased atmospheric carbon dioxide (CO₂) of700 µmol mol^–1 and increased air temperature of+ 4C were examined in Lolium perenne L. cv. Vigor, growingin semi-controlled greenhouses. Leaf growth, segmental elongationrates (SER), water relations, cell wall (tensiometric) extensibility(%P) and epidermal cell lengths (ECL) were measured in expandingleaves in spring and summer. In elevated CO₂, shoot dry weight (SDW) increased in mid-summer.In both seasons, SDW decreased in elevated air temperatureswith this reduction being greater in summer as compared to spring.Specific leaf area (SLA) decreased in elevated CO₂ and in CO₂ temperature in both seasons. In spring, increased leaf extensionand SER in elevated CO₂ were linked with increased ECL, %P andfinal leaf size whilst in summer all were reduced. In high temperature,leaf extension, SER, %P and final leaf size were reduced inboth seasons. In elevated CO₂ temperature, leaf extension,SER, %P, and ECL increased in spring, but final leaf size remainedunaltered, whilst in summer all decreased. Mid-morning waterpotential did not differ with CO₂ or temperature treatments.Leaf turgor pressure increased in elevated CO₂ in spring andremained similar to the control in summer whilst solute potentialdecreased in spring and increased in summer. Contrasting seasonalgrowth responses of L. perenne in response to elevated CO₂ andtemperature suggests pasture management may change in the future.The grazing season may be prolonged, but whole season productivitymay become more variable than today. Key words: Lolium perenne, ryegrass, CO₂ and temperature, leaf extension, cell wall rheology     

Effect of Elevated CO2 on Stomatal Density and Distribution in a C4 Grass and a C3 Forb under Field Conditions

Two common tallgrass prairie species, Andropogon gerardii, thedominant C₄ grass in this North American grassland, and Salviapitcheri, a C₃ forb, were exposed to ambient and elevated (twiceambient) CO₂ within open-top chambers throughout the 1993 growingseason. After full canopy development, stomatal density on abaxialand adaxial surfaces, guard cell length and specific leaf mass(SLM; mg cm^-2) were determined for plants in the chambers aswell as in adjacent unchambered plots. Record high rainfallamounts during the 1993 growing season minimized water stressin these plants (leaf xylem pressure potential was usually >-1·5 MPa in A. gerardii) and also minimized differencesin water status among treatments. In A. gerardii, stomatal densitywas significantly higher (190 ± 7 mm^-2; mean ±s.e.) in plants grown outside of the chambers compared to plantsthat developed inside the ambient CO₂ chambers (161 ±5 mm^-2). Thus, there was a significant 'chamber effect' on stomataldensity. At elevated levels of CO₂, stomatal density was evenlower (P < 0·05; 121 ± 5 mm^-2). Most stomatawere on abaxial leaf surfaces in this grass, but the ratio ofadaxial to abaxial stomatal density was greater at elevatedlevels of CO₂. In S. pitcheri, stomatal density was also significantlylower when plants were grown in the open-top chambers (235 ±10 mm^-2 outside vs. 140 ± 6 mm^-2 in the ambient CO₂ chamber).However, stomatal density was greater at elevated CO₂ (218 ±12 mm^-2) compared to plants from the ambient CO₂ chamber. Theratio of stomata on adaxial vs. abaxial surfaces did not varysignificantly in this herb. Guard cell lengths were not significantlyaffected by growth in the chambers or by elevated CO₂ for eitherspecies. Growth within the chambers resulted in lower SLM inS. pitcheri, but CO₂ concentration had no effect. In A. gerardii,SLM was lower at elevated CO₂. These results indicate that stomataland leaf responses to elevated CO₂ are species specific, andreinforce the need to assess chamber effects along with treatmenteffects (CO₂) when using open-top chambers.Copyright 1994, 1999Academic Press Andropogon gerardii, elevated CO₂, Salvia pitcheri, stomatal density, tallgrass prairie     

The Growth and Development of Maize (Zea mays L.) at Five Temperatures

The objectives of this work were to measure growth and developmentrates over a range of temperatures and to identify processeswhich may limit vegetative yield of maize (Zea mays L.). Twosingle cross Corn Belt Dent maize hybrids were grown from sowingin a diurnal temperature regime of 16/6 °C day/night andin constant temperature environments of 16, 20, 24 and 28 °C.The 16/6 °C environment was close to the minimum for sustainedgrowth and 28 °C was near the optimum. Entire plants wereharvested at stages with 4, 6, 7 and 8 mature leaves in alltemperature treatments except 20 °C in which the final twoharvests were carried out at 9 and 10 mature leaves. Mean totalleaf number varied between 19.5 and 16.0 with the maximum occurringat 16/6 °C. Although harvests were carried out at comparableleaf numbers, and hence at similar developmental stages, thetime interval between sowing and harvest decreased considerablyas temperatures increased. The relative rates of dry weight and leaf area accumulationwith time increased with a Q₁₀ of 2.4 between 16 and 28 °C,while leaf appearance rate increased with a Q₁₀ of 2.9 overthe same range; both rates were highest at 28 °C. Althoughdry matter partitioning to the shoots increased with temperature,the area of individual leaves varied in a systematic patternwhich resulted in maximum leaf area, leaf area duration andconsequently dry weight being realized at 20 °C for anygiven stage of development. Zea mays, corn, low temperature stress, temperature response, growth, development     

Influence of Wind Speed on the Sensitivity of Ryegrass to SO2

The sensitivity of S23 Lolium perenne L. to 11 parts 10^–8SO₂ was investigated at two different wind speeds. At the higherwind speed of 25 m min^–8, SO₂ caused significant reductionsin leaf area, root/shoot ratio, and all dry weight fractionsmeasured. At the lower wind speed of 10 m min^–1 growthreductions were not found. The differences in sensitivity ofplants to SO₂ at different wind speeds is discussed in relationto boundary layer resistances of leaves. It is concluded thatthe sensitivity of a plant to a particular pollutant shouldno longer be measured in terms of only the concentration andlength of exposure.     

Development of the Fifth Leaf is Indicative for Whole Plant Performance at Low Temperature in Tomato

In the search for early-detectable selection criteria for growthat low temperature conditions in tomato, first the initiationand growth of individual leaves was analysed. Scanning electronmicroscopy revealed that the first four primordia had alreadydeveloped during the germination period at 25°C. The primordiumof the fifth leaf, however, was initiated after the transferof seedlings to the experimental conditions. The increase inlength of the first three leaves, and to a lesser extent ofthe fourth leaf, was considerably smaller in comparison withthat of later formed leaves. Moreover, the morphology of thefirst three to four leaves was deviant, whereas the others showedthe normal compound leaf architecture. All these results indicatedthat the fifth leaf was the earliest formed leaf with growthcharacteristics that might reflect the growth potential of thewhole plant. Development of the fifth leaf was tested as a marker for wholeplant growth. At three temperature, 18, 15 and 12°C, growthresponses of the fifth leaf were similar to that of whole plantsin four tomato genotypes: Line A, Line B, Premier and MXXIV-13.Significant differences in relative growth rate of dry weightof whole plants and fifth leaves (RGR_W)and of leaf area of thefifth leaves (RGR_LA between two fast growing and two slow growinggenotypes were found. No genotype by temperature interactionfor RGR_W and RGR_LA was found, indicating that the effect oftemperature decrease was similar for the four genotypes. The structure of the mature fifth leaf of one fast and one slowgrowing genotype, Line A and MXXIV-13, was analysed. For bothgenotypes, leaves were small and thick at low temperature, 12°C.The total number of epidermis and palisade parenchyma cellsper leaf was smaller but the size of the cells developed at12°C was larger than at 18°C. Consequently, the slowgrowth at 12°C was due to a low rate of cell division. Atboth temperatures, the fifth leaf to MXXIV-13 was smaller comparedto that of line A. Since the size of the cells were similar,the smaller leaf size was due to lower number of leaf cells. The results confirm the suitability of the growth, especiallyexpressed as RGR_LA , of the fifth leaf as a nondestructive marketfor vegetative development of tomato at low temperature. Growthdifferences between genotypes were mainly reflected by differencesin cell number of leaves, which might be correlated with geneticallydetermined differences in cell number of leaf primordia.Copyright1993, 1999 Academic Press Lycopersicon esculentum Mill. genotypes, plant growth, selection criteria, low temperature, leaf initiation, leaf development, RGR, leaf structure, cell expansion