Photosynthesis by White Clover Leaves in Mixed Clover/Ryegrass Swards

Measurements of rates of net photosynthesis were made on singleBlanca white clover leaves on plants taken from a field-grown,mixed clover/perennial ryegrass sward during two regrowth periods. Net photosynthesis fell by 20 per cent in the first measurementperiod as leaf area index increased and the grass componentof the crop flowered, but did not change significantly in thesecond measurement period during which the grass remained vegetative. Leaves which had been artificially protected from shading inthe sward did not have significantly different photosyntheticcapacities from leaves in the undisturbed sward, even in thefirst measurement period. As leaf area index and sward height increased, successive cloverpetioles were longer, keeping the newly expanded leaves nearthe top of the sward where they received full light. It is suggestedthat it is this which allows successive clover leaves, unlikethose of vegetative grasses, to attain a high photosyntheticcapacity throughout a growth period. Trifolium repens, Lolium perenne, Photosynthetic capacity, shading, growth     

The Effect of Shading During Vegetative and Reproductive Growth on the Photosynthetic Capacity of Leaves in a Grass Sward

The decline in photosynthetic capacity of successive newly expandedleaves, which occurs as a vegetative grass sward increases inleaf area, was prevented in a field sward of S24 perennial ryegrass(Lolium perenne L.) by protecting tillers from shading by theirneighbours. This adds support for the view that the declineis caused by the shading of leaves during their expansion. However, in a flowering sward where there is no such decline,the photosynthetic capacity of leaves was not reduced by shadingthem with a plastic mesh during expansion, although it was bypegging down tillers so that they developed at the bottom ofthe sward. Lolium perenne, perennial rye-grass, photosynthetic capacity, growth, shading     

Effect of Flowering on the Photosynthetic Capacity of Ryegrass Leaves Grown With and Without Natural Shading

The photosynthetic capacity of newly expanded leaves of vernalizedor non-vernalized plants of S24 perennial ryegrass (Lolium perenneL.), grown in long or short photoperiods, was measured in twoexperiments. In the first, leaves were protected from shadingduring development, while in the second, the natural shade ofneighbouring tillers in a sward was allowed. In the first experiment there was little effect of vernalization,day length or flowering, and leaves in all treatments had photosyntheticrates at 250 W m^–2 of between 28 and 32 mg CO₂ dm^–2h_–1.In the second experiment the photosynthetic rate of successiveleaves fell as sward leaf area increased. This downward trendwas reversed, however, in flowering tillers in the vernalizedlong-day treatment, while in the other treatments, which didnot flower, photosynthetic capacity continued to fall. It isconcluded that the leaves of reproductive tillers have highphotosynthetic capacities because stem extension carries themto the top of the canopy where they are well illuminated duringexpansion. Lolium perenneL, ryegrass, photosynthetic capacity, flowering, shading, vernalization     

The Effects of Shading and Cutting Treatments on the Photosynthetic Rate of Ryegrass Leaves

As the leaf area of miniature simulated swards of S24 perennialreygrass (Lolium perenne L.) grown in controlled environmentincreased, successive leaves which expanded on the main stemof a sample plant within the sward had progressively lower photosyntheticcapacities. When the plants surrounding the sample plant were cut, the photosyntheticcapacity of the next leaf to expand was increased, whether ornot other leaves on the sample plant had been removed. Whenall fully expanded leaves on the sample plant were removed butthe surrounding plants left uncut, the photosynthetic capacityof successive leaves on the sample plant continued to decreaseas in uncut swards. These results support the view that the changes in photosyntheticcapacity in grass swards are consequences of changes in thelight climate of the developing leaf rather than of changesin demand for assimilates or growth substance supply. The declinein photosynthetic capactiy was contributed to by both stomataland residual diffusion resistances.     

The Effect of Temperature on Photosynthesis of Ryegrass and White Clover Leaves

The rate of photosynthesis of leaves of perennial ryegrass (Loliumperenne L.) and white clover (Trifollum pratense L.) grown atdifferent temperatures was measured at a range of temperatures.There was a small effect of the temperature at which a leafhad grown on its photosynthetic rate, but a large effect ofmeasurement temperature, especially in bright light, where photosyntheticrates at 15°C were about twice those at 5°C. It appearsthat temperature could affect sward photosynthesis in the field.Ryegrass and clover had similar photosynthetic rates which respondedsimilarly to temperature. Lolium perenne L., ryegrass, Trifolium pratense L., white clover, photosynthesis, temperature, irradiance     

The photosynthesis of ryegrass leaves grown in a simulated sward

Plants were taken from simulated swards of perennial ryegrass (Lolium perenne) grown in a controlled environment and the rates of photosynthesis of the youngest fully expanded leaves, and the second and third youngest leaves on the same tillers were measured. The youngest leaves had the highest rates and the third the lowest, with the second leaves intermediate. The rate of photosynthesis in bright light of successive youngest expanded leaves decreased as the swards increased in leaf area, but did not when plants were grown so that the main stem was not shaded. When plants were grown at different densities and the photosynthetic rates of leaves of a particular ontogenetic rank were measured, it was found that leaves on plants from higher densities had lower rates of photosynthesis. Also leaves on plants grown in bright light had higher photosynthetic rates than those on plants grown in dim light. It is concluded that the decline in the photosynthetic capacity of successive leaves in a rapidly growing simulated sward is due to the intense shading to which they are subjected during their expansion.     

Effect of Nitrogen Supply on the Photosynthetic Performance of Leaves from Coffee Plants Exposed to Bright Light

Although Coffea arabica L. grows naturally in shaded habitats,it can be cultivated under high light intensity, but not withoutsevere photoinhibition mainly during the period of transferfrom the nursery into the field. The present work examines someof the changes in the photosynthetic performance induced byexposure to high light and the possibility of using enhancednitrogen levels to overcome photoinhibition. For that purpose,young plants of Coffea arabica L. (cv. Catuai) grown in a shadedgreenhouse were treated with 0, 1 and 2 mmol of nitrogen and4 weeks later exposed to full solar irradiation, outside. Visible damage due to exposure to full sunlight appeared within2 d in all plants, resulting in a reduced photosynthetic leafarea and drastic shedding of leaves in the unfertilized plants.These effects were considerably less in plants with the highestN dose. After 130 d of exposure, there was 100% mortality inplants receiving no extra nitrogen, compared with 30% in theplants treated with 2 mmol nitrogen. Photosynthesis rates, leafconductance and transpiration presented minimum values after4 d of light stress. Large changes in the photosynthetic capacity(measured at high CO₂ concentration and high light intensity),quantum efficiency and fluorescence yield (F_v/F_m) indicate thatnet photosynthesis rate in the air had been reduced by bothstomatal closure and by changes at the photochemical level.All indicators show that N-fertilized plants were less affectedby photoinhibition. Key words: Coffee plant, nitrogen, photoinhibition, photosynthesis     

The Effect of Nitrogenous Fertilizer on the Photosynthesis and Growth of White Clover/Perennial Ryegrass Swards

The application of nitrogenous fertilizer in March to a whiteclover (cv. Blanca) and perennial ryegrass (cv. S23) sward resultedin a rapid suppression of the clover, relative to clover ina treatment given no added nitrogen. Thereafter, the cloverin both treatments grew more rapidly than the grass and itsproportion of the total leaf area in the mixture increased,as the leaf area index rose to 8. After a second applicationof N in early July, clover was not suppressed to the same extentas in the first growth period. Overall, the photosynthetic capacities of newly expanded cloverlaminae were similar in the two treatments. Clover laminae hadhigher photosynthetic capacities than grass, even in the grass-dominant+ N treatment. Lamina area, petiole length, and the number of live leaves perstolon were similar in the two treatments, indicating that thedifferences in total leaf area were due to the presence of fewerstolon growing points in the + N treatment. Trifolium repens L., white clover, Lolium perenne L., perennial ryegrass, nitrogen, leaf area index, photosynthesis, growth     

Microclimate, Canopy Structure and Photosynthesis in Canopies of Three Contrasting Temperate Forage Grasses: III. Canopy Photosynthesis, Individual Leaf Photosynthesis and the Distribution of Current Assimilate

The rates of canopy and individual leaf photosynthesis and ¹⁴Cdistribution for three temperate forage grasses Lolium perennecv. S24, L. perenne cv. Reveille and Festuc'a arundinacea cv.SI70 were determined in the field during a summer growth period.Canopy photosynthesis declined as the growth period progressed,reflecting a decline in the photosynthetic capacity of successiveyoungest fully expanded leaves. The decline in the maximum photosyntheticcapacity of the canopies was correlated with a decline in theirquantum efficiencies at low irradiance. Changes in canopy structureresulted in changes in canopy net photosynthesis and dark respiration.No clear relationships between changes in the environment andchanges in canopy net photosynthesis and dark respiration wereestablished. The relative distributions of ¹⁴C in the shootsof the varieties gave a good indication of the amount of drymatter per ground area in the varieties.     

Competition between grass and clover in spring as affected by nitrogen fertiliser

Detailed measurements of irrigated ryegrass/white clover swards growing without interruption, with or without nitrogen fertiliser in spring, showed that the relative growth rate of clover was as great as that of grass, in the + N sward, and considerably greater than grass, in the – N sward. Clover leaves were not overtopped by grass leaves. Indeed, in both swards, clover had a greater proportion of its leaf lamina area in the upper, well lit, layers of the canopy than grass did. Consequently, clover had a greater mean rate of leaf photosynthesis in situ in the sward than grass. Clover's advantage in photosynthetic rate per leaf area was offset to some extent by its smaller ratio of leaf area to total above-ground dry weight than grass. The consequences of these results for our understanding of competition between grass and clover in mixed swards are discussed.     

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     

Summer survival of leaves in a soft-leaved shrub (Phlomis fruticosa L., Labiatae) under Mediterranean field conditions: avoidance of photoinhibitory damage through decreased chlorophyll contents

Photosynthetic pigments and relative water content of youngleaves of P. fruticosa decreased considerably with the onsetof the summer dry period and stabilized at low values for thelast two summer months, while leaf growth was arrested. Correspondingdecreases in photochemical efficiency of photosystem II, asjudged by chlorophyll fluorescence measurements of predarkenedleaves, were, however, negligible. Following the first autumnheavy rains, growth was restored and photosynthetic pigmentsand relative water content increased to the pre-drought values.The results indicate that the reduction of chlorophylls doesnot result from severe photoinhibitory damage but, instead,it may be an adaptive response against the adverse conditionsof the Mediterranean summer. Some photosynthetic and photoprotective characteristics of P.fruticosa leaves at two stages of their development, i.e. atthe severely dehydrated state with arrested growth during latesummer and after their revival following the first heavy autumnrains were compared. Apart from the chlorophyll loss, the photonyield of O₂ evolution and the photosynthetic capacity at saturatedCO₂ were considerably suppressed during the summer, indicatingthat the extremely low net photosynthetic rates observed inthe field were the combined result of stomatal and mesophylllimitations. Epoxidation state was low at midday during thesummer, indicating an active, photodissipative xanthophyll cycle.Although zeaxanthin content did not increase at midday afterthe rains, the potential of the cycle was maintained in therevived leaves, as judged by the high concentrations of thecycle components. After the rains, the activities of the anti-oxidantenzymes (superoxide dismutase, ascorbate peroxidase) remainedrelatively unchanged on a chlorophyll basis, but increased whenexpressed on a leaf surface area or protein basis. It may beconcluded that P. fruticosa leaves avoid severe photoinhibitoryand oxidative damage during the long, warm, dry and sunny Mediterraneansummer by reducing light harvesting and electron flow capacity,whilst maintaining an adequate photoprotective ability. Thepreservation of a remarkable photodissipative and anti-oxidativepotential after the rains may be related to the low predictabilityof precipitation even during the rainy winter. Key words: Chl loss, anti-oxidant enzymes, xanthophyll cycle, photosynthesis, water stress     

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     

A Vertical Gradient of the Chloroplast Abundance among Leaves of Chenopodium album

The abundances of chloroplasts in leaves on the main stems ofChenopodium album at different height levels were investigatedin relation to the photosynthetic capacity and light environmentof the leaves. (1) The number of chloroplasts per mesophyllcell decreased with descending position of leaves, except foryoung developing leaves at the top of plants that had smallerchloroplast numbers per cell than matured leaves beneath them.Contents of chlorophyll and ribulose-1,5-bisphosphate carboxylase/oxygenaseper leaf area that were highest in the topmost young leavesand decreased with decreasing height level indicate that thereis a vertical gradient of chloroplast abundance per leaf areadecreasing from the top of the leaf canopy with depth. (2) Light-saturatingrate of photosynthetic oxygen evolution per leaf area of maturedleaves decreased more steeply with decreasing leaf positionthan the chloroplast number per cell. Gradients of chlorophylland the enzyme protein contents were also steeper than thatof the chloroplast number. Loss of photosynthesis in lower leavesis, therefore, ascribed partly to loss of whole chloroplastsand partly to reduced photosynthetic capacities of the remainingchloroplasts. (3) The chloroplast number per cell in newly expandedsecond leaves was comparable to those in leaves that have developedat later stages of the plant growth but decreased graduallyduring leaf senescence both in the dark and light. The formationof the vertical gradient of chloroplast abundance is, therefore,ascribed to loss of whole chloroplasts during senescence ofleaves. (4) Irradiance a leaf receives decreased sharply fromthe top of the canopy with depth. The physiological or ecophysiologicalsignificance of the vertical distribution of chloroplasts amongleaves was discussed taking light environments of leaves intoconsideration. (Received July 31, 1995; Accepted October 20, 1995)     

Control of the Acclimation of Photosynthesis to Light and Temperature in Relation to Partitioning of Photosynthate in Developing Soybean Leaves

Photosynthetic acclimation was examined by exposing third trifoliolateleaves of soybeans to air temperatures of 20 to 30°C andphotosynthetic photon flux densities (PPFD) of 150 to 950µmolphotons m^–2 s^–1 for the last 3 d before they reachedmaximum area. In some cases the environment of the third leafwas controlled separately from that of the rest of the plant.Photosynthesis, respiration and dry mass accumulation were determinedunder the treatment conditions, and photosynthetic capacity,and dry mass and protein content were determined at full expansion.Photosynthetic capacity, the light-saturated rate of net carbondioxide exchange at 25°C and 34 Pa external partial pressureof carbon dioxide, could be modified between 21 and 35 µmolCO₂ m^–2 s^–1 by environmental changes after leaveshad become exporters of photosynthate. Protein per unit leafmass did not differ between treatments, and photosynthetic capacityincreased with leaf mass per unit area. Photosynthetic capacityof third leaves was affected by the PPFD incident on those leaves,but not by the PPFD on other leaves on the plant. Photosyntheticcapacity of third leaves was affected by the temperature ofthe rest of the plant, but not by the temperature of the thirdleaves. Photosynthetic capacity was linearly related to carbondioxide exchange rate in the growth regimes, but not to daytimePPFD. At high PPFD, and at 25 and 30°C, mass accumulationwas about 28% of the mass of photosynthate produced. At lowerPPFD, and at 20°C, larger percentages of the photosynthateproduced accumulated as dry mass. The results suggest that photosynthatesupply is an important factor controlling leaf structural growthand, consequently, photosynthetic acclimation to light and temperature. Key words: Glycine max (L.) Merr., photosynthesis, temperature acclimation, light acclimation, photosynthate partitioning     

Photosynthesis, Water Relations, Temperature and Canopy Structure as Factors Influencing the Growth of Sainfoin (Onobrychis viciifolia Scop.) and Lucerne (Medicago sativa L.)

Measurements of the growth of sainfoin and lucerne were madein the field after cutting on 31 May 1977. Sainfoin reacheda total above-ground dry weight of 408 g m^–2 over thegrowing period of 48 days compared with 598 g m^–2 in lucerne.Final leaf area indices (LAIs) were 2.8 in sainfoin and 6.1in lucerne. The specific leaf areas (SLAs) for sainfoin wereapproximately half those of lucerne throughout the regrowthperiod. The maximum rates of leaf appearance were 0.12 leavesper day in sainfoin and 0.85 leaves per day in lucerne. Themaximum mean rate of plant extension growth for lucerne of 2.12mm h^–1 occurred during the night, whereas, in sainfointhe maximum rate of 1.72 mm h^–1 occurred during the day. Measurements of extinction coefficients for PAR ranged from0.45 to 0.89 in sainfoin and from 0 42 to 0.57 in lucerne. Asthe lucerne crop increased in size leaf water potentials andsolute potentials became more negative. In sainfoin leaf waterpotentials remained remarkably high throughout the growth period,solute potentials decreased and turgor potentials increased.The stomatal conductances of the two species were similar. The photosynthetic capacities and rates of dark respirationper unit leaf area in both species were similar. The rate ofcanopy ‘gross’ photosynthesis at 295 W m^–2was always greater in lucerne than in sainfoin. This was largelya matter of differences between the species in LAI, althoughat higher LAIs the more erect structure of lucerne leads toa better utilization of photosynthetically active radiation. Onobrychis vicifolia Scop, sainfoin, Medicago sativa L., lucerne, photosynthesis, water relations, temperature, canopy structure     

Ochna pulchra Hook: Leaf Growth and Development Related to Photosynthetic Activity

Ochna pulchra Hook. is a deciduous broad-leaved tree in theMixed Bushveld vegetation of the Northern Transvaal. The growthand development of leaves taken from trees in the field werestudied from a stage shortly before bud break, in late spring,until they were fully expanded and at the peak of photosyntheticactivity. Leaf area was measured by photographing the leaf against a transparentmm² grid. Finally a constant relationship between leaf area(A) and the linear dimension of length (L) and breadth (B) wasestablished: A = b x LB, where coefficient b = 0.72. Transverse sections of the lamina of the youngest leaves showeda five-layered plate meristem with a few functional conductingelements in the midrib. During further leaf development, celldivision was followed by means of autoradiography using [³H]thymidine.It was most active during the week after bud break. Leaf cell increment following on cell division made the majorcontribution to leaf growth resulting in a lamina that was atleast 90% expanded 4 weeks after bud break. The histologicalchanges accompanying cell division were observed using lightand electron microscopy. Even in late stages of leaf development mature and differentiatingstomata occurred together, limited to the abaxial epidermisand the midrib. Scanning electron microscopy showed stomataldistribution, their increasing density and gradual opening.The structure of these sunken stomata could reduce the outwarddiffusion of water vapour and increase the diffusion resistanceto carbon dioxide. Carbon assimilation rates of the developing leaves were measuredusing an IRGA (infra-red gas analyser) and their chlorophyllvalues were calculated. Photosynthesis was first measured amonth after bud break when the leaves were fully expanded, over50 % of the stomata exposed and leaf mesophyll tissue differentiatedwith mature chloroplasts. Net photosynthetic rates and chlorophyllvalues peaked 1 month later. Ochna pulchra Hook., photosynthesis, leaf development, leaf area, stomata, chlorophyll, savanna     

Mutual shading and the photosynthetic capacity of exposed leaves of field grown soybeans

Light-saturated photosynthetic rates at air levels of carbon dioxide were measured about weekly in upper canopy leaves of two soybean cultivars grown at stand densities of 40 and 100 plants per square meter. Early in the season, when leaf area indices differed between stand densities, plants of both cultivars grown at high stand density had photosynthetic rates which averaged 23% lower than plants at low stand density. Later in the season, when there were no differences in leaf area index between stand densities, there were no differences in photosynthetic rates in the cultivar Kent, but rate differences of about 14% persisted in the cultivar Williams. In Williams mainstem leaves emerged into full sunlight later in their development at high than at low stand density. In both cultivars the oldest fully exposed leaves were photosynthetically immature for much of the season, as higher rates could be achieved by lower leaves which were shaded in situ. The results identify shading of young developing leaves and photosynthetic immaturity of fully exposed leaves as factors limiting canopy photosynthesis in soybeans, and indicate cultivar differences in how much high stand density reduces photosynthetic capacity.     

Reactions of birch leaves to changes in light during early ontogeny: comparison between in vivo and in vitro techniques to measure carbon uptake

Trends in several photosynthetic parameters and their responseto changed growth light were followed for 15 d in leaves ofyoung birch saplings using a rapid-response gas exchange measuringequipment. These in vivo measurements were compared to biochemicalassays that were made from the same leaves after the gas exchangestudies. The measurements were made on leaves that were selectedprior to the study and were at that time of similar age. Forthe first 7 d the photosynthetic parameters were followed fromthe growth conditions of moderate light (200 µmol m^–2s^–1; referred to as controls later in the text). On day7 some of the saplings were transferred to grow either underhigh (450 µmol m^–2 s^–1; referred to as highlight plants) or low (75 µmol m^–2 s^–1; referredto as low light plants) light and the capability of the preselectedleaves for acclimation was followed for 6 d. For comparison,at the end of the experiment the measurements were made on bothcontrols and on young leaves that had developed under high andlow light. Generally the in vivo measured rate of CO₂ uptake (gross photosynthesis)both at 310 ppm CO₂ and 2000 ppm CO₂ corresponded very wellto the biochemically determined CO₂ fixation capacity in vitroafter rapid extraction (measured as the initial and total activityof Rubisco, respectively). However, if the flux of CO₂ intothe chloroplasts was limited by the closure of the stomata,as was the case of the high light plants, then the in vitromeasured Rubisco activity was greater than the in vivo measuredCO₂ uptake. V_max, calculated from the mesophyll conductanceat 1% O₂, exceeded the initial activity of Rubisco (assayedat saturating RuBP and CO₂) constantly by 60%. The catalyticactivity of Rubisco in birch leaves was overall very low, evenwhen calculated from the total activity of Rubisco (K_cat 0.63–1.18 s^–1), when compared to herbaceous C₃ species. Signs of light acclimation were not observed in most of thephotosynthetic parameters and in chloroplast structure whenmature birch leaves were subjected to changes in growth lightfor 6 d. However, the change of the growth light either to highor low light caused day-to-day fluctuations in most of the measuredphotosynthetic parameters and in the case of the high lightplants signs of photoinhibition and photodestruction were alsoobserved (decrease in the amount of chlorophyll and increasein chlorophyll a/b ratio). As a result of these fluctuationsthese plants achieved a new and lower steady-state conditionbetween the light and dark reactions, as judged from the molarratio of RuBP to Rubisco binding site. Key words: Acclimation, photosynthesis, light, Rubisco, birch     

Growth and Photosynthesis in the First Leaf of Barley The Effect of Time of Application of Nitrogen

Using Proctor barley grown in sand under controlled-environmentconditions it was shown that when application of nitrogen, asnitrate, was delayed beyond day 4, at which time the first leafwas beginning to unfold, absolute and relative growth-ratesof seedlings were reduced so that the young plants were significantlysmaller. Delay in nitrate application led to reduction in length,breadth, area, and dry weight of the first leaf, and also toa lower photosynthetic activity on day 8, as measured by infra-redgas analysis. Measurement of the uptake of ¹⁴CO² by first leaves showed thatapplication of nitrate on days 2 or 4 led to high rates of fixationof carbon over the period days 8–12, whereas applicationon days 6 or 8 led to a substantially lower maximum rate offixation which was maintained for a shorter period. When nitratewas applied on day 8 total fixation of carbon over the perioddays 7–14 was only 60 per cent of that for leaves on plantsfor which nitrate was supplied on day 2. When amounts of carbon fixed were compared on a leaf dry-weightbasis, maximum values were found to be similar for all treatments,suggesting that the differences in fixation per leaf resultmainly from the effect of treatment on leaf area. For all treatmentsit was confirmed that a decline in photosynthetic activity occurredby day 14. This was not correlated with photosynthetic activityin the second leaf, nor with emergence of the third leaf. Analysis of different parts of the lamina of the first leafshowed all to be affected by the timing of the supply, bothin growth and in photosynthetic activity. Leaf dry weight didnot increase after day 8 for any treatment, yet when nitratewas applied on day 8 plants showed a fourfold increase in photosyntheticrate. The significance of this in relation to carboxylationand other resistances in photosynthesis is discussed.