EFFECTS OF CANOPY POSITION AND IRRADIANCE ON THE LEAF PHYSIOLOGY AND MORPHOLOGY OF PENTACLETHRA MACROLOBA (MIMOSACEAE)

Pentaclethra macroloba (Willd.) Kuntze (Mimosaceae) is a dominant late-successional tree species in the Atlantic lowland forests of Costa Rica. Leaves of P. macroloba from three heights in the forest canopy were compared with leaves of seedlings grown in controlled environment chambers under four different irradiance levels. Changes in leaf characteristics along the canopy gradient paralleled changes resulting from the light gradient under controlled conditions. The effect of light or canopy position on light-saturated photosynthesis was small, with maximum photosynthesis increasing from 5 to 6.5 μmol m^−-2 s^−-1 from understory to canopy. Both chamber grown and field leaves showed large adjustments in photosynthetic efficiency at low light via reductions in dark respiration rates and increases in apparent quantum yields. Light saturation of all leaves occurred at or below 500 μmol m^−-2 s^−-1. Leaf thickness, specific leaf weight, and stomatal density increased to a greater extent than saturated photosynthesis with higher irradiance during growth or height in the canopy. As a result, there was a poor correspondence between leaf thickness and light-saturated photosynthesis on an area basis. It is concluded that Pentaclethra macroloba possesses the characteristics of a typical shade-tolerant species.     

Acclimation of photosynthesis to lightflecks in tomato leaves: interaction with progressive shading in a growing canopy

Plants in natural environments are often exposed to fluctuations in light intensity, and leaf‐level acclimation to light may be affected by those fluctuations. Concurrently, leaves acclimated to a given light climate can become progressively shaded as new leaves emerge and grow above them. Acclimation to shade alters characteristics such as photosynthetic capacity. To investigate the interaction of fluctuating light and progressive shading, we exposed three‐week old tomato (Solanum lycopersicum ) plants to either lightflecks or constant light intensities. Lightflecks of 20 s length and 1000 μmol m^?2 s^?1 peak intensity were applied every 5 min for 16 h per day, for 3 weeks. Lightfleck and constant light treatments received identical daily light sums (15.2 mol m^?2 day^?1). Photosynthesis was monitored in leaves 2 and 4 (counting from the bottom) during canopy development throughout the experiment. Several dynamic and steady‐state characteristics of photosynthesis became enhanced by fluctuating light when leaves were partially shaded by the upper canopy, but much less so when they were fully exposed to lightflecks. This was the case for CO₂‐saturated photosynthesis rates in leaves 2 and 4 growing under lightflecks 14 days into the treatment period. Also, leaf 2 of plants in the lightfleck treatment showed significantly faster rates of photosynthetic induction when exposed to a stepwise change in light intensity on day 15. As the plants grew larger and these leaves became increasingly shaded, acclimation of leaf‐level photosynthesis to lightflecks disappeared. These results highlight continuous acclimation of leaf photosynthesis to changing light conditions inside developing canopies.     

Simultaneous gas exchange and fluorescence measurements indicate differences in the response of sunflower,bean and maize to water stress

Gas exchange and fluorescence measurements of attached leaves of water stressed bean, sunflower and maize plants were carried out at two light intensities (250 mol quanta m^-2s^-1 and 850 mol quanta m^-2s^-1). Besides the restriction of transpiration and CO₂ uptake, the dissipation of excess light energy was clearly reflected in the light and dark reactions of photosynthesis under stress conditions. Bean and maize plants preferentially use non-photochemical quenching for light energy dissipation. In sunflower plants, excess light energy gave rise to photochemical quenching. Autoradiography of leaves after photosynthesis in ¹⁴CO₂ demonstrated the occurrence of leaf patchiness in sunflower and maize but not in bean. The contribution of CO₂ recycling within the leaves to energy dissipation was investigated by studies in 2.5% oxygen to suppress photorespiration. The participation of different energy dissipating mechanisms to quanta comsumption on agriculturally relevant species is discussed.Abbreviations F_o minimal fluorescence - F_m maximal fluorescence - F_p peak fluorescence - g leaf conductance - P_N net CO₂ uptake - q_N coefficient of non-photochemical quenching - q_P coefficient of photochemical quenching     

Photooxidative damage to the photosynthetic apparatus during chilling

Short periods of chilling in the presence of light (up to 6 h: 1°C; 270 W/m²) decreased the subsequent apparent photosynthesis and chlorophyll fluorescence of leaf discs of Cucumis sativus L. cv. Kleine Groene Scherpe. The extent of the injury depended on the duration of the chilling pretreatment. After 6 h the subsequent apparent photosynthesis even reached a negative value, and it increased only slightly during the next 2 1/2 h. The decrease of apparent photosynthesis was not a consequence of increased dark respiration but was of photooxidative origin since the presence of both light and oxygen was required. Preincubation in the light for 2 h at 20°C sensitized leaf discs to subsequent photooxidation during chilling. Prompt and delayed chlorophyll fluorescence decreased simultaneously after chilling and light treatment. The corresponding decrease of photosynthesis and chlorophyll fluorescence is discussed in relation to primary photooxidative damage to the photosystems in the chloroplast membrane.     

LIGHT,TEMPERATURE AND SALINITY EFFECTS ON GROWTH,LEAF ANATOMY AND PHOTOSYNTHESIS OF DISTICHLIS SPICATA (L.) GREENE

The effects of light, temperature, and salinity on growth, net CO₂ exchange and leaf anatomy of Distichlis spicata were investigated in controlled environment chambers. When plants were grown at low light, growth rates were significantly reduced by high substrate salinity or low temperature. However, when plants were grown at high light, growth rates were not significantly affected by temperature or salinity. The capacity for high light to overcome depressed growth at high salinity cannot be explained completely by rates of net photosynthesis, since high salinity caused decreases in net photosynthesis at all environmental conditions. This salinity-induced decrease in net photosynthesis was caused largely by stomatal closure, although plants grown at low temperature and low light showed significant increases in internal leaf resistance to CO₂ exchange. Increased salinity resulted in generally thicker leaves with lower stomatal density but no significant differences in the ratio of mesophyll cell surface area to leaf area. Salinity and light during growth did not significantly affect rates of dark respiration. The mechanisms by which Distichlis spicata tolerates salt appear to be closely coulpled to the utilization of light energy. Salt-induced leaf succulence is of questionable importance to gas exchange at high salinity in this C₄ species.     

Screening a cucumber crop during leaf area development reduces yield

In order to reduce heat energy consumption in greenhouse cucumber production, (transparent) screens may be used also during the day, particularly in the early growth phase when high temperatures are required to achieve rapid leaf area development. However, energy savings must be optimised against light reduction‐induced yield loss. For this reason, two experiments were conducted to quantify the effect on photosynthesis and growth of screening cucumber plants during their early growth phase, and on yield in the following generative phase. Screening with different light transmission coefficients was simulated using shading nets. Shading the plants during the first 5 weeks under Central European winter conditions reduced the leaf area by 0.40% per 1% reduction in photosynthetic active radiation (PAR). Moreover, potential leaf net photosynthesis decreased by 0.46% per 1% PAR reduction. A major impact was that the leaf dry matter content, leaf starch content and leaf sugar content of shaded plants diminished significantly. In the course of the following 2 weeks under full light, the leaf photosynthesis of the plants previously shaded recovered fully and the leaf area index rose to 3.3 m² m^?2, considered sufficient for optimal crop photosynthesis. The yield from plants previously shaded diminished slightly as early as from the first harvest week on. These yield losses increased further over the next few weeks, measuring approximately 0.8 kg m^?2 per 1 mol m^?2 day^?1 PAR reduction in the early growth phase. The effect of PAR on plant growth was proportional when relating the PAR integral over the entire experimental period to the total yield and to the total dry matter production, respectively.     

RESPONSE OF PROROCENTRUM MARIAE-LEBOURIAE (DINOPHYCEAE) TO LIGHT OF DIFFERENT SPECTRAL QUALITIES AND IRRADIANCES: GROWTH AND PIGMENTATION1

Growth and pigment concentrations of the, estuarine dinoflagellate, Prorocentrum mariae-lebouriae (Parke and Ballantine) comb. nov., were measured in cultures grown in white, blue, green and red radiation at three different irradiances. White irradiances (400–800 nm) were 13.4, 4.0 and 1.8 W · m^?2 with photon flux densities of 58.7 ± 3.5, 17.4 ± 0.6 and 7.8 ± 0.3 μM quanta · m^?2· s^?1, respectively. All other spectral qualities had the same photon flux densities. Concentrations of chlorophyll a and chlorophyll c were inversely related to irradiance. A decrease of 7- to 8-fold in photon flux density resulted in a 2-fold increase in chlorophyll a and c and a 1.6- to 2.4-fold increase in both peridinin and total carotenoid concentrations. Cells grown in green light contained 22 to 32% more peridinin per cell and exhibited 10 to 16% higher peridinin to chlorophyll a ratios than cells grown in white light. Growth decreased as a function of irradiance in white, green and red light grown cells but was the same at all blue light irradiances. Maximum growth rates occurred at 8 μM quanta · m^?2· s^?1 in blue light, while in red and white light maximum growth rates occurred at considerably higher photon flux densities (24 to 32 μM quanta · m^?2· s^?1). The fastest growth rates occurred in blue and red radiation. White radiation producing maximum growth was only as effective as red and blue light when the photon flux density in either the red or blue portion of the white light spectrum was equivalent to that of a red or of blue light treatment which produced maximum growth rates. These differences in growth and pigmentation indicate that P. mariae-lebouriae responds to the spectral quality under which it is grown.     

Light and growth temperature alter carbon isotope discrimination and estimated bundle sheath leakiness in C<Subscript>4</Subscript> grasses and dicots

We combined measurements of short-term (during gas exchange) and long-term (from plant dry matter) carbon isotope discrimination to estimate CO₂ leakiness from bundle sheath cells in six C₄ species (three grasses and three dicots) as a function of leaf insertion level, growth temperature and short-term irradiance. The two methods for determining leakiness yielded similar results (P > 0.05) for all species except Setaria macrostachya, which may be explained by the leaf of this species not being accommodating to gas exchange. Leaf insertion level had no effect on leakiness. At the highest growth temperature (36°C) leakiness was lower than at the two lower growth temperatures (16°C and 26°C), between which no differences in leakiness were apparent. Higher irradiance decreased leakiness in three species, while it had no significant effect on the others (there was an opposite trend in two species). The inverse response to increasing irradiance was most marked in the two NAD-ME dicots (both Amaranthus species), which both showed almost 50% leakiness at low light (300 μmol quanta m⁻² s⁻¹) compared to about 30% at high light (1,600 μmol quanta m⁻² s⁻¹). NADP-ME subtype grasses had lower leakiness than NAD-ME dicots. Although there were exceptions, particularly in the effect of irradiance on leakiness in Sorghum and Boerhavia, we conclude that conditions favourable to C₄ photosynthesis (high temperature and high light) lead to a reduction in leakiness.     

Improvement of acclimatization of micropropagated grapevine: Photosynthetic competence and carbon allocation

Grapevine plantlets multiplied in vitro were acclimatized at 40 or 90 μmol m⁻² s⁻¹ photon flux density for 12 or 16 h per day, respectively. In the high-light regime a decrease in total chlorophyll and an increase in chlorophyll a/chlorophyll b ratio occurred. However, at high-light intensity lower photosynthetic capacities and higher apparent photosynthesis were measured than at the low-light regime. In leaves expanded during acclimatization, the light compensation point was higher in plantlets under high-light while quantum yield was higher in low-light conditions. High-light also gave rise to an increase in carbohydrate concentration. As a whole, the results suggest that high-light increases carbon assimilation and growth although with a low investment in the photosynthetic apparatus. This revised version was published online in June 2006 with corrections to the Cover Date.     

LIGHT ABSORPTION BY PLANTS AND ITS IMPLICATIONS FOR PHOTOSYNTHESIS

The preceding account has attempted to examine the interactions between light absorption and photosynthesis, with reference to both unicellular and multicellular terrestrial and aquatic plants. There are, however, some notable plant groups to which no direct reference has been made, e.g. mosses, liverworts and lichens. Although many have similar optical properties to terrestrial vascular plants (Gates, 1980) and apparently similar photosynthetic responses (see Green & Snelgar, 1982; Kershaw, 1984) they may possess subtle, as yet unknown differences. For instance, the lichen thallus has a high surface reflectance although the transmittance is virtually zero (Gates, 1980; Osborne, unpublished results). It is envisaged, however, that differences in optical properties between species will reflect differences in degree not kind. Although not all variation in photosynthesis is due to differences in light absorption a number of accounts suggest that this is a contributing factor. Variations in leaf absorptance have been found to account for most of the variation in leaf photosynthesis at low J_is (see Ehleringer & Björkman, 1978a; Osborne & Garrett, 1983). There is, however, little direct experimental evidence on light absorption and photosynthesis in either microalgal species or aquatic macrophytes. We also do not know over what range of incident photon flux densities photosynthesis is determined largely by changes in light absorption. Plants growing under natural conditions also experience large diurnal and seasonal fluctuations in J_i, unlike species grown under laboratory conditions. The occurrence of transitory peaks in J_i tends to overshadow the fact that the average J_i is often lower than the J₁ required to saturate photosynthesis, i.e. 1500–2000 μmol m^-2 s^-1, depending on the growth treatment. Using the data of Monteith (1977) and I W m²= 5 μmol m^-2 s^-1, and with photosynthetically active radiation 50% of total solar radiation, the daily mean value for Britain is approximately 450 μmol m^-2 s^-1, with a maximum in June of 1000μmol m^-2 s^-1 and a minimum during the winter of 75 μmol m^-2 s^-1. Such values could be even lower on shaded understory leaves and considerably lower for aquatic species. Based on average values of net photosynthesis for a terrestrial plant leaf, light saturation would only be expected in June while for most of the year the average values would lie largely on the light-limited portion of the photosynthesis light response curve. Although the daily average values in tropical climates may be higher during the winter months, they are remarkably similar throughout the world for the respective summers in the northern and southern hemispheres, because the increased daylength at high latitudes compensates for the lower J_is. The expected lower dark respiration rates during the winter may also partially offset the effects of a lower light level. There is therefore a trade-off between high J_is for a short period of time against a lower J_i for a longer period of time. We might expect different photosynthetic responses to these two very different conditions. Importantly, a low J_i with a long daylength may enable a plant to photosynthesize at or near its maximum photon efficiency for most of the day. Although the response of the plant to fluctuations in J_i is complicated because it is affected by the previous environmental conditions, this may indicate that light absorption has a much greater significance under natural conditions, particularly for perennial species. The bias in many laboratories towards research on terrestrial vascular plants also tends to ignore the fact that a number of multicellular and unicellular aquatic species survive in very low light environments. Furthermore, the direct extrapolation of photosynthetic responses from measurements on single leaves to those of whole plants is clearly erroneous. Although this is obvious, many physiological ecologists have attributed all manner of things to the photosynthetic responses of ‘primary’ leaves. Most researchers have ignored problems associated with composite plant tissues and internal light gradients. Clearly caution is required in interpreting the photosynthesis light-response curve of multicellular tissues based on biochemical features alone. Also, the importance of cell structure on light absorption and photosynthesis has generally been ignored and attributed solely to the effects of structural features on CO₂ diffusion. In doing so the work of two or three generations of plant physiologists has been ignored. Haberlandt (1914) at the turn of the century probably first implicated the role of cell structure in leaf optics, and Heath (1970) stressed that in order to completely understand the role of light in photosynthesis we need to know the flux incident on the chloroplast itself. Even this suggestion may need modification because of the capacity of the internal chloroplast membranes for scattering light. It is worth emphasizing the importance of light gradients within tissues and their role in regulating photosynthesis, particularly at light saturation. Measurements of light gradients are fraught with problems because of experimental difficulties and the majority (few) are based on reflectance and transmittance measurements. Seyfried & Fukshansky (1983) have shown that light incident on the lower surface of a Cucurbita cotyledon produced a larger light gradient than light incident from above, indicating the importance of the spatial arrangement of the tissues with respect to the light source. Also, light incident on the lower surface of leaves of Picea sitchensis was less ‘effective’ in photosynthesis than light from above (Leverenz & Jarvis, 1979). Clearly, two tissues could have the same gross absorptance but different photosynthetic rates because of differences in the internal light environment. Fisher & Fisher (1983) have recently found asymmetries in the light distribution within leaves, which they related to asymmetries in photosynthetic products due to differences in solar elevation. Such modifications in light distribution could be important for a number of solar-tracking species. Changes in light absorption are brought about by a whole gamut of physiological, morphological and behavioural responses which serve to optimize the amount of light absorbed. Perhaps the simplest way of regulating the amount of light absorbed is by restricting growth either to particular times of the year or to conditions when the light climate is favourable. We are still largely ignorant of many details of these modifications. In particular, differences in tissue structure such as the size and number of vacuoles or the effects of organelles on the scattering component of the internal light environment of photosynthetic tissues are not understood. A better understanding of the interaction of light with plants in aquatic systems is also required. It is unfortunate that light-absorptance measurements are not routinely made in photosynthetic studies, and this is quite clearly a neglected area of study. That these measurements are not made is even more surprising, since techniques have been available for over sixty years (Ulbricht, 1920). Absorptance measurements are of particular importance in the photosynthetic adaptation of microalgae, where only a small proportion of the incident photon flux density is absorbed. For multicellular species more detailed information is required on internal light gradients and their variability. Light-absorptance measurements are also important in any study relating kinetic data on CO₂ fixation to in vivo photosynthesis, especially when there are large variations in the morphology and structure of the photosynthetic organ.     

Variation in leaf respiration in relation to growth and photosynthesis of Lolium

Six Lolium genotypes with contrasting apparent photorespiration and CO_a compensation concentration, [C0₂]_c, were compared for net photosynthesis, dark respiration, leaf starch accumulation, rate of leaf expansion and shoot regrowth. Plants were grown in day/night temperatures of 15/10 and 25/20 ^oC. There were significant (P < 0–05) differences between the genotypes in all these parameters. At 25/20 ^oC apparent photorespiration was correlated with [CO₂]_c. Correlation coefficients, pooled from both temperature regimes, revealed that genotypes with high rates of net photosynthesis accumulated more leaf starch during light periods than genotypes with slow photosynthesis, but rates of leaf expansion and dry matter increase were only correlated, negatively, with dark respiration. Apparent photorespiration was negatively correlated with dark respiration. These findings suggest that attributes related to photorespiration such as [CO₂]_c and O₂ uptake from CO₂-free air in the light are unlikely to be useful selection criteria for growth of C₃ grasses, that net photosynthesis was probably not limiting growth and that maintenance respiration may have been an important determinant of genotypic differences in growth rate. Selections for slow and fast rates of dark respiration of mature leaves were therefore made at 8 and at 25 ^oC from within two different populations of L. perenne, S.23. This characteristic showed repeatabilities (broad-sense heritability) of from 0–41 to o-66. Six independent comparisons of simulated swards of the slow- and fast-respiring selections were made under periodic cutting regimes, either in a growth room at 25 ^oC or in a glasshouse from August to May. Growth of all plots of slow-respiring genotypes was consistently more rapid than that of the fast-respiring, at 25 ^oC in the growth room, and during autumn and spring in the glasshouse. There was no difference in winter growth. The implications of these results for the use of gas exchange measurements as selection criteria in plant breeding programmes are discussed.     

THE INFLUENCE OF WATER STRESS ON STEM AND LEAF PHOTOSYNTHESIS IN GLYCINE MAX AND SPARTEUM JUNCEUM (LEGUMINOSAE)

Stem and leaf photosynthesis were measured in Glycine max var. essex (soybean) and Sparteum junceum (Spanish broom). The significance of stem photosynthesis to whole plant growth was evaluated by blocking stem photosynthesis with black straw sections. The growth of S. junceum was reduced by 18% when black straws were used in comparison to clear straws. The whole plant growth of G. max was not influenced by blocking the stem carbon contribution. Mean midday leaf photosynthesis was 12 μmol CO₂ m^–2 s^–1 and 17 μmol CO₂ m^–2 s^–1 for G. max and 5. junceum, respectively. Mean midday stem photosynthesis of S. junceum was 6.5 μmol CO₂ m^–2 s^–1; however, positive net photosynthesis did not occur in G. max stems. Water stress caused a proportionally greater decrease in leaf photosynthesis compared to that of stems during diurnal cycles of photosynthesis in S. junceum. As a result the contribution to canopy carbon gain by stem photosynthesis increased from 38% to 48% of the total plant carbon gain under reduced water availability.     

Seasonal relationships between leaf nitrogen content (photosynthetic capacity) and leaf canopy light exposure in peach (Prunus persica)

Abstract Field gas exchange measurements on intact peach (Prunus persica (L.) Batsch) leaves indicate that leaf nitrogen content (N_L) and leaf weight per unit leaf area (W_a) are highly correlated with CO₂ assimilation rate (A) and mesophyll conductance (g_m). Therefore, N_L and W_a were used to study seasonal relationships between leaf carboxylation capacity and natural light exposure in tree canopies. From mid-season onwards, N_L and W_a were linearly correlated with light exposure expressed as the amount of time during a clear day that a leaf was exposed to a photosynthetic photon flux density (Q) of ≥ 100 μmol m^?2 s^?1. The data support the hypothesis that whole-tree photosynthesis is optimized by partitioning of photosynthetic capacity among leaves in deciduous tree canopies with respect to natural light exposure.     

Modelling photosynthetic photon flux density and maximum potential gross photosynthesis

Irradiance data software developed by the NREL Solar Radiation Laboratory (Simple Model of Atmospheric Radiative Transfer of Sunshine, SMARTS) has been used for modelling photosynthesis. Spectra and total irradiance were expressed in terms of quanta [mol m⁻² s⁻¹, photosynthetic photon flux density, PPFD (400–700 nm)]. Using the SMARTS software it is possible to (1) calculate the solar spectrum for a planar surface for any given solar elevation angle, allowing for the attenuating effects of the atmosphere on extraterrestrial irradiance at each wavelength in the 400–700 nm range and for the thickness of atmosphere the light must pass through during the course of a day, (2) calculate PPFD vs. solar time for any latitude and date and (3) estimate total daily irradiance for any latitude and date and hence calculate the total photon irradiance for a whole year or for a growing season. Models of photosynthetic activity vs. PPFD are discussed. Gross photosynthesis (P _g) vs. photosynthetic photon flux density (PPFD) (P _g vs. I) characteristics of single leaves compared to that of a canopy of leaves are different. It is shown that that the optimum irradiance for a leaf (I_opt) is the half-saturation irradiance for a battery of leaves in series. A C₃ plant, with leaves having an optimum photosynthetic rate at 700 μmol m⁻² s⁻¹ PPFD, was used as a realistic worked example. The model gives good estimates of gross photosynthesis (P _g) for a given date and latitude. Seasonal and annual estimates of P _g can be made. Taking cloudiness into account, the model predicts maximum P _g rates of about 10 g(C) m⁻² d⁻¹, which is close to the maximum reported P _g experimental measurements.     

PHOTOSYNTHESIS AND CARBON ALLOCATION IN TIPULARIA DISCOLOR (ORCHIDACEAE), A WINTERGREEN UNDERSTORY HERB

Seasonal patterns of photosynthesis and carbon allocation were determined for Tipularia discolor, a summer-deciduous wintergreen orchid of the southeastern United States, to assess the effects of environmental conditions and leaf age on carbon acquisition and allocation patterns. There was no shift in the optimum temperature for photosynthesis (T_opt) on a seasonal basis and T_opt (≈26 C) was at least 10 C higher than daily maximum air temperature during most of the growing season. Lack of photosynthetic adjustment in Tipularia to seasonal fluctuations in temperature and light suggested that the photosynthetic characteristics of this wintergreen were more similar to those of spring ephemerals than to those of evergreens and summer-active herbs. The decline in photosynthetic capacity during the winter growing season for Tipularia, largely due to leaf age effects, gradually reduced net photosynthetic rates in the field despite more favorable light and temperature conditions. Photosynthesis in the field was primarily limited by environmental conditions in early- and mid-season and by photosynthetic capacity in late-season. A ¹⁴CO₂ labelling experiment demonstrated that patterns of carbon allocation to vegetative structures were affected by the season of photosynthetic carbon fixation, whereas reproductive structures received 21% of the recovered labelled carbon regardless of the period of labelling. Carbon acquired and stored during all periods of the growing season was used to produce new vegetative and reproductive structures.     

Response of total night-time respiration to differences in total daily photosynthesis for leaves in a Quercus rubra L. canopy: implications for modelling canopy CO2 exchange

Measurements of photosynthesis and respiration were made on leaves in summer in a Quercus rubra L. canopy at approximately hourly intervals throughout 5 days and nights. Leaves were selected in the upper canopy in fully sunlit conditions (upper) and in the lower canopy (lower). In addition, leaves in the upper canopy were shaded (upper shaded) to decrease photosynthesis rates. The data were used to test the hypothesis that total night‐time respiration is dependent on total photosynthesis during the previous day and that the response is mediated through changes in storage in carbohydrate pools. Measurements were made on clear sunny days with similar solar irradiance and air temperature, except for the last day when temperature, especially at night, was lower than that for the previous days. Maximum rates of photosynthesis in the upper leaves (18.7 μmol m^?2 s^?1) were approximately four times higher than those in the lower leaves (4.3 μmol m^?2 s^?1) and maximum photosynthesis rates in the upper shaded leaves (8.0 μmol m^?2 s^?1) were about half those in the upper leaves. There was a strong linear relationship between total night‐time respiration and total photosynthesis during the previous day when rates of respiration were normalized to a fixed temperature of 20°C, removing the effects of temperature from this relationship. Measurements of specific leaf area, nitrogen and chlorophyll concentration and calculations of the maximum rate of carboxylation activity, V_cmax, were not significantly different between upper and upper shaded leaves 5 days after the shading treatment was started. There were small, but significant decreases in the rate of apparent maximum electron transport at saturating irradiance, J_max (P>0.05), and light use efficiency, ? (P<0.05), for upper shaded leaves compared with those for upper leaves. This suggests that the duration of shading in the experiment was sufficient to initiate changes in the electron transport, but not the carboxylation processes of photosynthesis. Support for the hypothesis was provided from analysis of soluble sugar and starch concentrations in leaves. Respiration rates in the upper shaded leaves were lower than those expected from a relationship between respiration and soluble sugar concentration for fully exposed upper and lower leaves. However, there was no similar difference in starch concentrations. This suggests that shading for the duration of several days did not affect sugar concentrations but reduced starch concentrations in leaves, leading to lower rates of respiration at night. A model was used to quantify the significance of the findings on estimated canopy CO₂ exchange for the full growing season. Introducing respiration as a function of total photosynthesis on the previous day resulted in a decrease in growing season night‐time respiration by 23% compared with the value when respiration was held constant. This highlights the need for a process‐based approach linking respiration to photosynthesis when modelling long‐term carbon exchange in forest ecosystems.     

Analysis of fluorescence transients of DCMU-treated leaves of Triticum species to provide estimates of the densities of photosystem II reaction centres

The fluorescence of the chlorophyll associated with photosystem II was studied in seedling and flag leaves of Triticum species. Seedling leaves of the diploid species T. urartu had higher values of t (the normalised area over the fluorescence induction curve of DCMU treated leaves) than those of the other species studied which included hexaploid T. aestivum. However this difference was not evident when leaves were grown in a low light intensity (40 µmol quanta of photosynthetically active radiation m^–2 s^–1). The smaller total number of chlorophyll molecules per photosystem II reaction centre (chl/RCII) in T. urartu (177) as compared with the other species (mean 234) was deduced from the observed differences in t. As a consequence of its lower chl/RCII, despite slightly lower chlorophyll content (mg m^–2), T. urartu had a greater density of reaction centres than the other species (2880 cf 2230 nmol m^–2 of leaf). Consistent with the lower chl/RCII of T. urartu, it had a higher chlorophyll a/b ratio than the other genotypes. Seedling leaves of T. urartu had higher light saturated rates of photosynthesis than those of the other species, when grown at high light, a difference associated with reaction centre density.In flag leaves, when the complications due to variable development and senescence patterns were eliminated, t of the diploid species including T. urartu was lower than that of T. aestivum. The lower apparent chl/RCII of T. urartu arose partly because the molar extinction coefficient of the chlorophyll in the leaves of T. urartu was greater than in T. aestivum. However, the density of PS II reaction centres was slightly lower for the diploid species studied because their chlorophyll contents were lower than the hexaploids.The validity of the method for estimating chl/RCII from fluorescence transients is discussed. The possibility is considered that the difference in apparent chl/RCII of flag and seedling leaves of R. urartu as compared to the other five genotypes is a consequence of its different adaptive response to the spectral quality of the light.     

Adaptations of the Photosynthetic Mechanism of Sugar Maple (Acer saccharum) Seedlings Grown in Various Light Intensities

Sugar maple (Acer saccharum Marsh.) seedlings were grown in a nursery for three years in 13, 25, 45 and 100 per cent of full daylight. During the third year of growth, the rates of their apparent photosynthesis and respiration were measured periodically with an infra-red gas analyzer at various light intensities and normal CO₂ concentration. In addition, the rates of apparent photosynthesis of a single attached leaf of the same seedlings were measured at saturating light intensity, hut varying CO₂ concentrations. An increase in the light intensity in which seedlings were grown had no effect on their height or mean leaf area, hut resulted in thicker leaves, an increase in the total leaf area per seedling due to an increase in the number of leaves, an increase in the dry weight especially of roots and a decrease in the chlorophyll content of leaves. Throughout the growing season seedlings grown in full daylight, as compared with those grown in lower light intensities, had the lowest rates of apparent photosynthesis measured at standard conditions (21,600 lux light intensity and 300 ul/l of CO₂), when this was expressed per unit leaf area, hut the highest rates on a per seedling basis. Thus dry matter production attained at the end of the growing season correlated positively with the photosynthetic rate per seedling, but not per unit leaf area. The rates of apparent photosynthesis of seedlings grown at lower light intensities were more responsive to changes in light intensity or CO₂ concentration than those of seedlings grown in full daylight intensity.     

Effect of light interception by non-photosynthetic organs on canopy photosynthetic production

A canopy photosynthesis model was derived on the assumption that the light diminution within a canopy is caused by both leaves and non-photosynthetic organs. The light diminution by leaves and that by non-photosynthetic organs were taken into account separately in the Lambert-Beer equation of light extinction. The light flux density on the leaf surface at each depth was evaluated from the leaf's share of light. The light flux density on the leaf surface thus obtained was incorporated into the Monsi-Saeki model of canopy photosynthesis. The proposed model was applied for estimating gross canopy photosynthesis in a 19-year-oldLarix leptolepis plantation where 38% of the light diminution was due to non-photosynthetic organs. The daily canopy photosynthesis on one summer day calculated using the present model was about 22% less than that calculated by the conventional Monsi-Saeki model, in which light interception by non-photosynthetic organs is neglected. The degree of such reduction in canopy photosynthesis through shading by non-photosynthetic organs was assessed in relation to parameters affecting light extinction, leaf photosynthetic characteristics, and light regime above the canopy.