Importance of the green color,absorption gradient,and spectral absorption of chloroplasts for the radiative energy balance of leaves

Terrestrial green plants absorb photosynthetically active radiation (PAR; 400–700 nm) but do not absorb photons evenly across the PAR waveband. The spectral absorbance of photosystems and chloroplasts is lowest for green light, which occurs within the highest irradiance waveband of direct solar radiation. We demonstrate a close relationship between this phenomenon and the safe and efficient utilization of direct solar radiation in simple biophysiological models. The effects of spectral absorptance on the photon and irradiance absorption processes are evaluated using the spectra of direct and diffuse solar radiation. The radiation absorption of a leaf arises as a consequence of the absorption of chloroplasts. The photon absorption of chloroplasts is strongly dependent on the distribution of pigment concentrations and their absorbance spectra. While chloroplast movements in response to light are important mechanisms controlling PAR absorption, they are not effective for green light because chloroplasts have the lowest spectral absorptance in the waveband. With the development of palisade tissue, the incident photons per total palisade cell surface area and the absorbed photons per chloroplast decrease. The spectral absorbance of carotenoids is effective in eliminating shortwave PAR (<520 nm), which contains much of the surplus energy that is not used for photosynthesis and is dissipated as heat. The PAR absorptance of a whole leaf shows no substantial difference based on the spectra of direct or diffuse solar radiation. However, most of the near infrared radiation is unabsorbed and heat stress is greatly reduced. The incident solar radiation is too strong to be utilized for photosynthesis under the current CO₂ concentration in the terrestrial environment. Therefore, the photon absorption of a whole leaf is efficiently regulated by photosynthetic pigments with low spectral absorptance in the highest irradiance waveband and through a combination of pigment density distribution and leaf anatomical structures.     

Transmission properties of an oak canopy in relation to photoperception

Abstract The spectra of incoming daylight and shadelight in a mature oak woodland were measured at intervals during the canopy cycle, and mean transmittance spectra were derived. Transmittance was spectrally neutral at ca. 0.55 during the light phase but, following leaf emergence, transmittance of PAR (400–700 nm) fell to ca. 0.1. Simultaneously, the red : far-red transmittance ratio fell to circa 0.6. Both showed little change during the summer and autumn until senescence, indicating that the optical properties of the canopy were surprisingly stable. There was no evidence that cloud cover influenced mean canopy transmittance, although transient sunflecks introduced great variability which, in combination with sampling bias, might explain previous contradictory reports. The red : far-red fluence rate ratio in the woodland showed a temporary increase in late summer, a result of a small increase in the red : far-red ratio of incoming daylight during this period. Reflectance and transmittance spectra and pigment content of sun and shade leaves were measured. Leaf transmittance spectra showed changes correlated with those of the canopy, and were related to changes in pigment content.     

The composition and function of thylakoid membranes from pea plants grown under white or green light with or without far-red light

Pea plants ( Pisum sativum L. ev. Greenfeast) were grown for 2 to 3 weeks in while (˜ 50 μmol photons m⁻² s⁻¹; 400–700 nm) or green (˜ 30 μmol photons m⁻² s ⁻¹ 400–700 nm) light (16 h day/8 h night), with or without far-red light. Supplementary far-red light decreased leaf area and increased internodal length in both white and green light, demonstrating that phytochrome influenced leaf size and plant growth. However, there was no effect of far-red light on chlorophyll a /chlorophyll b ratios, chlorophyll-protein composition, the stoichiometry of electron transport complexes or photosynthetic function of isolated thylakoids. These results suggest that phytochrome is ineffective in modulating the composition and function of thylakoids in pea plants grown at low irradiance. One possible explanation of the ineffectiveness of phytochrome on thylakoids is discussed in terms of the drastic attenuation of red relative to far-red light in green tissue.     

Scaling CO2-photosynthesis relationships from the leaf to the canopy

Responses of individual leaves to short-term changes in CO₂ partial pressure have been relatively well studied. Whole-plant and plant community responses to elevated CO₂ are less well understood and scaling up from leaves to canopies will be complicated if feedbacks at the small scale differ from feedbacks at the large scale. Mathematical models of leaf, canopy, and ecosystem processes are important tools in the study of effects on plants and ecosystems of global environmental change, and in particular increasing atmospheric CO₂, and might be used to scale from leaves to canopies. Models are also important in assessing effects of the biosphere on the atmosphere. Presently, multilayer and big leaf models of canopy photosynthesis and energy exchange exist. Big leaf models — which are advocated here as being applicable to the evaluation of impacts of global change on the biosphere — simplify much of the underlying leaf-level physics, physiology, and biochemistry, yet can retain the important features of plant-environment interactions with respect to leaf CO₂ exchange processes and are able to make useful, quantitative predictions of canopy and community responses to environmental change. The basis of some big leaf models of photosynthesis, including a new model described herein, is that photosynthetic capacity and activity are scaled vertically within a canopy (by plants themselves) to match approximately the vertical profile of PPFD. The new big leaf model combines physically based models of leaf and canopy level transport processes with a biochemically based model of CO₂ assimilation. Predictions made by the model are consistent with canopy CO₂ exchange measurements, although a need exists for further testing of this and other canopy physiology models with independent measurements of canopy mass and energy exchange at the time scale of 1 h or less.Abbreviations LAI leaf area index - NIR near infrared (700–3000 nm) radiation - PAR photosynthetically active (400–700 nm) radiation - PI photosynthetic irradiance (400–700 nm) - PPFD photosynthetic photon flux area density (400–700 nm) - PS I Photosystem I - PS II Photosystem II - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuP₂ ribulose-1,5-bisphosphate     

Modelling environmental limits to light use efficiency for a canopy of two broad-leaved tree species with contrasting leaf habit

We used outputs from a model of canopy carbon uptake [Dungan et al. (2004) Functional Ecology 18: 34–42] and measurements of irradiance (PAR, 400–700 nm) intercepted by the canopy to investigate the effect of daily changes in environmental conditions on daily light use efficiency, ε, for a canopy comprising two broadleaved New Zealand tree species with contrasting leaf habit. Irradiance absorbed by the canopy was 93% of the incident irradiance, and seasonal changes in the proportion of this absorbed by leaves of each species was estimated with a detailed model of leaf area phenology. Over the year, ε for semi-deciduous wineberry (Aristotelia serrata) was 0.43 g C MJ-1 PAR, with maximum and minimum values of 0.80 g C MJ-1 PAR and 0.07 g C MJ-1 PAR in summer and winter respectively. In contrast annual ε was 0.60 g C MJ-1 PAR for winter deciduous fuchsia, with a maximum value of 0.92 g C MJ-1 PAR in spring. The most important environmental regulator of ε for both species was τ, atmospheric transmissivity. Maximum values for ε were estimated on days when τ ≈0.2, on cloudy days in mid-summer. Limits to photosynthesis from restricted root-zone water availability were also important, showing that drought limitations can restrict ε even at a field site with annual rainfall of 4800 mm. Environmental limits to photosynthesis and ε have been investigated for only a few canopy tree species. Uncertainty in models of the national carbon budget required for reporting purposes would be reduced by considering the environmental regulation of ε for a wider range of tree species.     

Photomorphogenesis and canopy dynamics. Phytochrome-mediated proximity perception accounts for the growth dynamics of canopies of Populus trichocarpa × deltoides'Beaupré'

The phytochrome family of signal-transducing photoreceptors provides plants with the capacity to perceive variations in the relative fluxes of red (R) and far-red (FR) radiation. This capacity has been proposed to be of ecological value in the perception of the proximity of neighbouring plants and the consequent induction of shade avoidance responses. The work reported here has evaluated this potential by determining quantitatively the effect of neighbour proximity on the growth of canopies of Populus trichocarpa×deltoides‘Beaupré’ trees, and relating the measured variables to the long-term vectoral radiation quality inside each canopy. The spectral distribution of radiation inside four canopies of Populus trichocarpa×deltoides‘Beaupre’ of different densities was monitored throughout the growing season. Spectral distributions inside the canopies were measured in 10° wedges at different heights and angles. The results are presented as PFD over 400–700 nm (PFD400–700) and PFD over 400–800 nm (PFD400–700). Results are also presented for the calculated phytochrome photoequilibrium (Pfr/P) and red:far-red ratio (R:FR). Data are presented as in-canopy angular and height profiles, and as diurnal and seasonal variations. PFD_400–700 and Pfr/P were found to be reduced inside each canopy, the reduction being greatest in the most dense canopy, and least in the most open canopy. At any height within each canopy, calculated Pfr/P decreased linearly with time throughout the growing season, until leaf senescence began. The reduction was greater in the denser canopies and was found to be similar for three consecutive field seasons. Linear relationships were found between plant stem growth rate, plant spacing and Pfr/P calculated from radiation propagated approximately horizontally within the canopies. The findings support the role of phytochrome in proximity perception in the natural environment and provide a quantitative basis for investigating the competitive interactions between plants growing in dense stands. The hypothesis is proposed that the dynamics of developing or regenerating canopies can be accounted for on the basis of phytochrome-mediated perception of the proximity of neighbouring plants.     

Light absorption by phytoplankton and chromophoric dissolved organic matter in the drainage basin and estuary of the Neuse River, North Carolina (U.S.A.)

1. We examined the absorption of solar radiation by phytoplankton and chromophoric dissolved organic matter (CDOM) taking into account riparian shading in the rivers, reservoirs, swamps of the Neuse River Estuary and its drainage basin. 2. In the streams, CDOM typically absorbed 55 and 64% of photons in the spectral range of 400–700 nm (photosynthetically active radiation, PAR) and 500–600 nm, respectively. The large proportion of photons absorbed by CDOM indicates high potential for abiotic photochemial reactions in the 500–600 nm region. 3. Despite the high concentration of nutrients, phytoplankton contributed little (2%) to the total absorption of PAR in the streams. Small (<30 m wide) streams typically received only 7% of incident PAR that impinged onto the more exposed reservoirs and estuary. Riparian shading and the low contribution of phytoplankton to the total absorption resulted in conditions where phytoplankton absorbed nearly two orders of magnitude less PAR in the streams than in the estuary and reservoirs. 4. The results indicated that riparian shading and non‐algal absorbing components can significantly restrict phytoplankton production in nutrient‐rich streams with a high concentration of CDOM flowing throughout forested catchments.     

不同水氮条件下水稻冠层反射光谱与植株含水率的定量关系

研究了不同土壤水氮条件下水稻 (Oryzasativa) 冠层光谱反射特征和植株水分状况的量化关系。结果表明, 水稻冠层近红外光谱反射率随土壤含水量的降低而降低, 短波红外光谱反射率随土壤含水量的降低而升高。相同土壤水分条件下, 高氮水稻的冠层含水率高于低氮水稻的冠层含水率 ;同一水分条件下, 高氮处理的可见光区和短波红外波段光谱反射率低于低氮处理, 近红外波段光谱反射率高于低氮处理。发现拔节后比值植被指数 (R810 /R460 ) 与水稻叶片含水率和植株含水率呈极显著的线性相关, 模型的检验误差 (RootmeansquareError, RMSE) 分别为 0.93和 1.5 0。表明比值植被指数R810 /R460 可以较好地监测不同生育期水稻叶片和植株含水率。     

Modification of leaf cytology and anatomy in Brassica napus grown under above ambient levels of supplemental UV-B radiation.

Plants exposed to natural solar radiation usually show acclimation responses on a daily and seasonal basis. Many of these responses are complex and modified by interactions with acclimation responses to other climatic factors. While changes in photosynthetically active radiation (PAR, 400-700 nm) are the driving force for many acclimation responses in plants, radiation outside the PAR range is also important. Recently, interest has increased in the potential role of UV-A (320-400 nm) and UV-B (280-320 nm) components of sunlight in plant developmental, physiological and daily acclimation processes. In order to explore the role of UV-B further, Brassica napus L. cv Paroll plants were grown to maturity under 13 kJ d(-1) of biologically effective ultraviolet-B radiation (UV-B(BE), 280-320 nm) plus 800 micromol photons m(-2) s(-1) photosynthetically active radiation (PAR, 400-700 nm) or PAR alone. Leaf anatomy and palisade cell structure were quantified using stereological techniques. The leaves of plants grown under UV-B radiation exhibited an increase in overall leaf width, although no change in leaf anatomy was discerned. Palisade cells in UV-B exposed leaves showed a significant decrease in chloroplast, mitochondrial, starch, and microbody volume density (Vv), while the vacuolar Vv increased compared to cells exposed to PAR only. In UV-B exposed leaves, there was an increase in the appressed and non-appressed thylakoid surface area density (Sv) within the chloroplasts. Since the relative proportion of appressed to non-appressed thylakoid surface area did not change, both thylakoid systems changed in concert with each other. Thylakoid stacks were broader and shorter in leaves subjected to UV-B. In general these responses were similar to those which occurred in plants moved from a high to low PAR environment and similar to mature plants exposed to 13 kJ d(-1) UV-B(BE) for only a short period of time. Although UV absorbing pigments increased by 21% in UV-B exposed leaves, there was no significant difference in chlorophyll a,b or carotenoid content compared to plants exposed to only PAR.     

Leaf area index, canopy structure and photosynthesis of red clover (Trifolium pratense L.)

Abstract. The influence of leaf age, total leaf area and its dispersion in space on canopy photosynthesis were studied using microswards of red clover ( Trifolium pratense L.) which were established in the greenhouse. Two varieties, Renova (flowering) and Molstad (non-flowering), were sown in separate plastic boxes at densities of 225, 400 and 625 plants per m².
Vertical distribution of photosynthetically active radiation (PAR), leaf area, leaf age and ¹⁴CO₂-fixation were determined periodically. Net photosynthesis and dark respiration of canopies were measured. Maximum photosynthetic capacity of individual leaves was measured on plants taken from the intact canopy or from plants where shading of the growing leaves had been prevented.
Net photosynthetic rate of canopies increased linearly with leaf area index (LAI) up to an LAI of 3.5 and then declined at higher LAI, independent of variety and sowing density. Below the optimum LAI, net photosynthesis depended mainly on interception of PAR. Decrease in canopy photosynthesis above the optimum LAI was due to a higher proportion of old leaves with decreased photosynthetic capacity, and not to an increase in respiring plant parts. It is concluded that LAI and position of leaf age categories in the canopy are more important than vertical distribution of leaf area in determining canopy photosynthesis of red clover.     

Characterization of a newly isolated freshwater Eustigmatophyte alga capable of utilizing far-red light as its sole light source

Oxygenic phototrophs typically utilize visible light (400–700 nm) to drive photosynthesis. However, a large fraction of the energy in sunlight is contained in the far-red region, which encompasses light beyond 700 nm. In nature, certain niche environments contain high levels of this far-red light due to filtering by other phototrophs, and in these environments, organisms with photosynthetic antenna systems adapted to absorbing far-red light are able to thrive. We used selective far-red light conditions to isolate such organisms in environmental samples. One cultured organism, the Eustigmatophyte alga Forest Park Isolate 5 (FP5), is able to absorb far-red light using a chlorophyll (Chl) a-containing antenna complex, and is able to grow under solely far-red light. Here we characterize the antenna system from this organism, which is able to shift the absorption of Chl a to >705 nm.     

THE CONCHOCELIS OF PORPHYRA HAITANENSIS (RHODOPHYTA) IS PROTECTED FROM HARMFUL UV RADIATION BY THE COVERING CALCAREOUS MATRIX1

Previous study has shown that Porphyra conchocelis is sensitive to high levels of PAR (400–700 nm) as well as ultraviolet radiation (UVR: 280–400 nm), resulting in high inhibition of photosynthesis. However, little is known about whether the inner covering layer of the shell, in which the conchocelis lives, may provide protection against solar UVR. Our study indicates that the covering calcareous matrix is about 0.06 mm thick, transmitting 63, 47, and 28% of PAR, ultraviolet radiation A (UVA: 315–400 nm), and ultraviolet radiation B (UVB: 280–315 nm), respectively. We used a shading layer that simulated the above transmissions, and the effective quantum yield of PSII and photosynthetic carbon fixation in the conchocelis increased to greater extents in the presence of UVA or UVB. Attenuation of UVA by 19% and UVB by 37% due to the shading layer increased the PSII yield by 44%–77% and photosynthetic carbon fixation by about 60%. Our study clearly shows that the photosynthetic machinery of Porphyra haitanensis T. J. Chang et B. F. Zheng conchocelis was efficiently protected from harmful UVR by the covering calcareous matrix.     

UV-B and PAR in a grass (Lolium perenne L.) canopy

The penetration of natural and artificial UV-B_BE (Biologically Effective UV-B, Caldwell 1971) and PAR (400–700 nm) in a grass canopy with increasing LAI was followed during 2 months. Overall, the transmission of UV-B_BE sunlight is significantly higher than of PAR sunlight. This is mainly due to the higher proportion of diffuse light in the UV-B. Under cloudy conditions no difference between UV-B_BE and PAR could be found. Sun angle and intensity of the radiation were less important in determining the penetration of light. Artificial light penetrates much more through the canopy, resulting in higher irradiation levels in the lower part of the canopy, but a lower UV-B_BE/PAR ratio (since UV-B transmittance of the leaves is lower). The UV-B_BE/PAR ratio reaching the leaves was influenced by LAI, sun angle, percent diffuse light and leaf angle. The large differences in UV-B_BE/PAR ratio per unit leaf area under natural and artificial light conditions are important in understanding the influence of UV-B on plants.     

Photosynthetic response of Nodularia spumigena to UV and photosynthetically active radiation depends on nutrient (N and P) availability

Biomass of N. spumigena is distributed within the dynamic photic zone that changes in both light quantity and quality. This study was designed to determine whether nutrient status can mitigate the negative impacts of experimental radiation treatments on the photosynthetic performance of N. spumigena. Cyanobacterial suspensions were exposed to radiation consisting of photosynthetically active radiation (PAR=400-700 nm), PAR+UV-A (=PA, 320-700 nm), and PAR+UV-A+UV-B (=PAB, 280-700 nm) under different nutrient media either replete with external dissolved nitrate (N) and orthophosphate (P; designated as +N/+P), replete with P only (-N/+P), or replete with N only (+N/-P). Under low PAR (75 micromol photons m(-2) s(-1)), nutrient status had no significant effect on the photosynthetic performance of N. spumigena in terms of rETRmax, alpha, and E(k). Nodularia spumigena was able to acclimate to high PAR (300 micromol photons m(-2) s(-1)), with a corresponding increase in rETRmax and E(k). The photosynthetic performance of N. spumigena cultured with supplemental nitrogen was more susceptible to experimental PAR irradiance. Under UVR, P-enrichment in the absence of additional external N (-N/+P) induced lower photoinhibition of photosynthesis compared with +N/-P cultures. However, the induction of NPQ may have provided PSII protection under P-deplete and PAR+UVR conditions. Because N. spumigena are able to fix nitrogen, access to available P can render them less susceptible to photoinhibition, effectively promoting blooms. Under a P-deficient condition, N. spumigena were more susceptible to radiation but were capable of photosynthetic recovery immediately after removal of radiation stress. In the presence of an internal P pool in the Baltic Sea, which may be seasonally available to the diazotrophic cyanobacteria, summer blooms of the resilient N. spumigena will persist.     

Some observations on the spectral distribution characteristics of short-wave radiation within Pinus radiata D. Don canopies

Abstract. Measurements of the photosynthetic photon flux density (400–700 nm) and of the spectral distribution of photon flux density across the 370–800 nm waveband, were made under both clear and overcast sky conditions above and at various positions within two Pinus radiata canopies of different stocking but similar leaf area indices. The spectra obtained for the daylight conditions (i.e. above forest canopy) were generally similar to those published previously. The spectra for shadelight within the forest canopy showed no blue peak which was characteristic of previously reported measurements which were restricted to the diffuse radiation component. There was almost neutral absorption within the 400–700 nm waveband, and typical lower attenuation in the 700–800 nm waveband. The blue: red ratio was largely unchanged by either canopy type or sky conditions and varied between 0.57 and 0.81. The red: far-red ratio in shadelight was between 0.22 and 0.41 under clear sky and between 0.68 and 0.95 under overcast sky conditions. Values for daylight were between 1.16–1.22. Calculated phytochrome photoequilibrium values in shadelight were approximately 0.35 under clear sky and 0.46 under overcast sky conditions. In each case there appeared to be no differences between the two canopies with respect to these minimum values.     

Effects of kaolin application on light absorption and distribution, radiation use efficiency and photosynthesis of almond and walnut canopies

BACKGROUND AND AIMS: Kaolin applied as a suspension to plant canopies forms a film on leaves that increases reflection and reduces absorption of light. Photosynthesis of individual leaves is decreased while the photosynthesis of the whole canopy remains unaffected or even increases. This may result from a better distribution of light within the canopy following kaolin application, but this explanation has not been tested. The objective of this work was to study the effects of kaolin application on light distribution and absorption within tree canopies and, ultimately, on canopy photosynthesis and radiation use efficiency. METHODS: Photosynthetically active radiation (PAR) incident on individual leaves within the canopy of almond (Prunus dulcis) and walnut (Juglans regia) trees was measured before and after kaolin application in order to study PAR distribution within the canopy. The PAR incident on, and reflected and transmitted by, the canopy was measured on the same day for kaolin-sprayed and control trees in order to calculate canopy PAR absorption. These data were then used to model canopy photosynthesis and radiation use efficiency by a simple method proposed in previous work, based on the photosynthetic response to incident PAR of a top-canopy leaf. KEY RESULTS: Kaolin increased incident PAR on surfaces of inner-canopy leaves, although there was an estimated 20 % loss in PAR reaching the photosynthetic apparatus, due to increased reflection. Assuming a 20 % loss of PAR, modelled photosynthesis and photosynthetic radiation use efficiency (PRUE) of kaolin-coated leaves decreased by only 6.3 %. This was due to (1) more beneficial PAR distribution within the kaolin-sprayed canopy, and (2) with decreasing PAR, leaf photosynthesis decreases less than proportionally, due to the curvature of the photosynthesis response-curve to PAR. The relatively small loss in canopy PRUE (per unit of incident PAR), coupled with the increased incident PAR on the leaf surface on inner-canopy leaves, resulted in an estimated increase in modelled photosynthesis of the canopy (+9 % in both walnut and almond). The small loss in PRUE (per unit of incident PAR) resulted in an increase in radiation use efficiency per unit of absorbed PAR, which more than compensated for the minor (7 %) reduction in canopy PAR absorption. CONCLUSIONS: The results explain the apparently contradictory findings in the literature of positive or no effects of kaolin applications on canopy photosynthesis and yield, despite the decrease in photosynthesis by individual leaves when measured at the same PAR.     

Motility and photosynthetic responses of the green microalga Tetraselmis subcordiformis to visible and UV light levels

To test the effects of photosynthetic active radiation (PAR, 400–700 nm) and ultraviolet radiation (UVR, 280–400 nm) on phototaxis and photosynthesis of free swimming microalgae, experiments were performed with Tetraselmis subcordiformis (Wille) Butcher under a solar simulator. In particular, we evaluated the effects of different PAR levels and radiation regimes (i.e., PAR only and PAR+UVR) on those two processes. We found that the cells preferred to move to a particular area (e.g., receiving 100 W m^?2 PAR) with little photochemical suppression or inhibition of carbon fixation. Adding UV-A to high PAR decreased its swimming capacity and photosynthetic capability, and further adding UV-B led to more inhibition. The suppression of the moving capability of T. subcordiformis was reversible but the cells exposed to PAR combined with UVR needed longer time intervals to recover their motility as compared with those irradiated only with PAR. Based on the above results, we postulate that in nature, the motile capability and photosynthesis of free swimming the green microalga might be impaired by enhanced solar UVR. On the other hand, the cells can reduce the damage caused by high irradiances (and even get the optimum light level for photosynthesis) by a behavioral swimming response.     

Estimating photosynthetic radiation use efficiency using incident light and photosynthesis of individual leaves

It has been theorized that photosynthetic radiation use efficiency (PhRUE) over the course of a day is constant for leaves throughout a canopy if leaf nitrogen content and photosynthetic properties are adapted to local light so that canopy photosynthesis over a day is optimized. To test this hypothesis, 'daily' photosynthesis of individual leaves of Solanum melongena plants was calculated from instantaneous rates of photosynthesis integrated over the daylight hours. Instantaneous photosynthesis was estimated from the photosynthetic responses to photosynthetically active radiation (PAR) and from the incident PAR measured on individual leaves during clear and overcast days. Plants were grown with either abundant or scarce N fertilization. Both net and gross daily photosynthesis of leaves were linearly related to daily incident PAR exposure of individual leaves, which implies constant PhRUE over a day throughout the canopy. The slope of these relationships (i.e. PhRUE) increased with N fertilization. When the relationship was calculated for hourly instead of daily periods, the regressions were curvilinear, implying that PhRUE changed with time of the day and incident radiation. Thus, linearity (i.e. constant PhRUE) was achieved only when data were integrated over the entire day. Using average PAR in place of instantaneous incident PAR increased the slope of the relationship between daily photosynthesis and incident PAR of individual leaves, and the regression became curvilinear. The slope of the relationship between daily gross photosynthesis and incident PAR of individual leaves increased for an overcast compared with a clear day, but the slope remained constant for net photosynthesis. This suggests that net PhRUE of all leaves (and thus of the whole canopy) may be constant when integrated over a day, not only when the incident PAR changes with depth in the canopy, but also when it varies on the same leaf owing to changes in daily incident PAR above the canopy. The slope of the relationship between daily net photosynthesis and incident PAR was also estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident PAR above the canopy, in place of that measured on individual leaves. The slope (i.e. net PhRUE) calculated in this simple way did not differ statistically from that calculated using data from individual leaves.     

Exposure to ultraviolet radiation delays photosynthetic recovery in Arctic kelp zoospores

Seasonal reproduction in some Arctic Laminariales coincides with increased UV-B radiation due to stratospheric ozone depletion and relatively high water temperatures during polar spring. To find out the capacity to cope with different spectral irradiance, the kinetics of photosynthetic recovery was investigated in zoospores of four Arctic species of the order Laminariales, the kelps Saccorhiza dermatodea, Alaria esculenta, Laminaria digitata, and Laminaria saccharina. The physiology of light harvesting, changes in photosynthetic efficiency and kinetics of photosynthetic recovery were measured by in vivo fluorescence changes of Photosystem II (PSII). Saturation irradiance of freshly released spores showed minimal I _k values (photon fluence rate where initial slope intersects horizontal asymptote of the curve) values ranging from 13 to 18 μmol photons m⁻² s⁻¹ among species collected at different depths, confirming that spores are low-light adapted. Exposure to different radiation spectra consisting of photosynthetically active radiation (PAR; 400–700 nm), PAR+UV-A radiation (UV-A; 320–400 nm), and PAR+ UV-A+UV-B radiation (UV-B; 280–320 nm) showed that the cumulative effects of increasing PAR fluence and the additional effect of UV-A and UV-B radiations on photoinhibition of photosynthesis are species specific. After long exposures, Laminaria saccharina was more sensitive to the different light treatments than the other three species investigated. Kinetics of recovery in zoospores showed a fast phase in S. dermatodea, which indicates a reduction of the photoprotective process while a slow phase in L. saccharina indicates recovery from severe photodamage. This first attempt to study photoinhibition and kinetics of recovery in zoospores showed that zoospores are the stage in the life history of seaweeds most susceptible to light stress and that ultraviolet radiation (UVR) effectively delays photosynthetic recovery. The viability of spores is important on the recruitment of the gametophytic and sporophytic life stages. The impact of UVR on the zoospores is related to the vertical depth distribution of the large sporophytes in the field.     

The ratio of transmitted near-infrared radiation to photosynthetically active radiation (PAR) increases in proportion to the adsorbed PAR in the canopy

The daily total photosynthetically active radiation (400??00?nm, PAR) and near-infrared radiation (700??000?nm, NIR) were measured in the understory beneath the canopy (PARt and NIRt) and above the canopy (PARi and NIRi) of a Japanese cool-temperate deciduous broad-leaved forest during the snow-free period (May to November). The integration of spectral radiation for NIR and that for PAR, and the daily integrations of instantaneous NIR and PAR, reduced the noises from the optical difference in spectrum and from canopy structure heterogeneity, sky condition and solar elevation. PARi/PARt was linearly related to NIRt/PARt (R2?=?0.96). The effect of cloudiness was negligible, because the fluctuation of NIRi/PARi was quite small regardless of season and weather conditions compared with the range of NIRt/PARt in the forest. The ratio of NIRt/PARt beneath the canopy was log-linearly related to the in situ leaf area index (LAI) with a wide range from 0 to 5.25 (R2?=?0.97). We conclude that seasonal changes in fAPAR (=?1???PARt/PARi) and LAI of a canopy can be estimated with high accuracy by transmitted NIRt and PARt beneath the canopy.