Basal leaf senescence in a sunflower (Helianthus annuus) canopy: responses to increased R/FR ratio

Senescence of lower leaves (LS) begins before anthesis in sunflower crop canopies. Using isolated field-grown sunflower plants, it has previously been shown that pre-anthesis LS is dependent on photosynthetic photon flux density (PPFD) and is hastened by increases in far-red light. We tested the hypothesis that increasing the red/far-red ratio (R/FR) perceived by basal leaves within canopies delays LS. To do this, light impinging on the lower surface of north-oriented 8th leaves (cotyledons=0) of crops with maximum leaf area indexes of 3.3 (Experiment 1) and 2.4 (Experiment 2) was enriched (+8.33 μmol m⁻² s⁻¹) with red light using light emitting diode (LED) panels. LED panels constructed with unlit LED or with green LED (PPFD slightly greater than the red LED panels, to compensate for lower efficiency) were used as controls. Compared with controls, additional R significantly ( P <0.05) increased R/FR perceived by the lower surface and significantly ( P <0.01) delayed LS. On average, leaf duration, as time between full expansion and a 70% diminution of chlorophyll content, was 5 days greater for leaves receiving extra red light (maximum observed LD=27 days). We conclude that an increase in the R/FR ratio can delay LS in crop canopies.     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model     

Modelling canopy CO2 fluxes: are ‘big‐leaf’ simplifications justified?

• 1 The ‘big‐leaf’ approach to calculating the carbon balance of plant canopies is evaluated for inclusion in the ETEMA model framework. This approach assumes that canopy carbon fluxes have the same relative responses to the environment as any single leaf, and that the scaling from leaf to canopy is therefore linear.
• 2 A series of model simulations was performed with two models of leaf photosynthesis, three distributions of canopy nitrogen, and two levels of canopy radiation detail. Leaf‐ and canopy‐level responses to light and nitrogen, both as instantaneous rates and daily integrals, are presented.
• 3 Observed leaf nitrogen contents of unshaded leaves are over 40% lower than the big‐leaf approach requires. Scaling from these leaves to the canopy using the big‐leaf approach may underestimate canopy photosynthesis by ~20%. A leaf photosynthesis model that treats within‐leaf light extinction displays characteristics that contradict the big‐leaf theory. Observed distributions of canopy nitrogen are closer to those required to optimize this model than the homogeneous model used in the big‐leaf approach.
• 4 It is theoretically consistent to use the big‐leaf approach with the homogeneous photosynthesis model to estimate canopy carbon fluxes if canopy nitrogen and leaf area are known and if the distribution of nitrogen is assumed optimal. However, real nitrogen profiles are not optimal for this photosynthesis model, and caution is necessary in using the big‐leaf approach to scale satellite estimates of leaf physiology to canopies. Accurate prediction of canopy carbon fluxes requires canopy nitrogen, leaf area, declining nitrogen with canopy depth, the heterogeneous model of leaf photosynthesis and the separation of sunlit and shaded leaves. The exact nitrogen profile is not critical, but realistic distributions can be predicted using a simple model of canopy nitrogen allocation.

     

Leaf canopy as a dynamic system: ecophysiology and optimality in leaf turnover

BACKGROUND AND AIMS: In a leaf canopy, there is a turnover of leaves; i.e. they are produced, senesce and fall. These processes determine the amount of leaf area in the canopy, which in turn determines canopy photosynthesis. The turnover rate of leaves is affected by environmental factors and is different among species. This mini-review discusses factors responsible for leaf dynamics in plant canopies, focusing on the role of nitrogen. SCOPE: Leaf production is supported by canopy photosynthesis that is determined by distribution of light and leaf nitrogen. Leaf nitrogen determines photosynthetic capacity. Nitrogen taken up from roots is allocated to new leaves. When leaves age or their light availability is lowered, part of the leaf nitrogen is resorbed. Resorbed nitrogen is re-utilized in new organs and the rest is lost with dead leaves. The sink-source balance is important in the regulation of leaf senescence. Several models have been proposed to predict response to environmental changes. A mathematical model that incorporated nitrogen use for photosynthesis explained well the variations in leaf lifespan within and between species. CONCLUSION: When leaf turnover is at a steady state, the ratio of biomass production to nitrogen uptake is equal to the ratio of litter fall to nitrogen loss, which is an inverse of the nitrogen concentration in dead leaves. Thus nitrogen concentration in dead leaves (nitrogen resorption proficiency) and nitrogen availability in the soil determine the rate of photosynthesis in the canopy. Dynamics of leaves are regulated so as to maximize carbon gain and resource-use efficiency of the plant.     

Effects of canopy shade on the lipid composition of soybean leaves

The effect of canopy shade on leaf lipid composition was examined in soybeans ( Glycine max cv. Young) grown under field conditions. Expanding leaves were tagged at 50, 58 and 65 days after planting (DAP) in plots with either a high (10 plants m⁻¹ row) or low (1 plant m⁻¹ row) plant density. At 92 DAP, light conditions ranged from a pho-tosynthetic photon flux density (PPFD) of 87% of full sun with a far-red/red (735 nm/645 nm) ratio of 0.9 at upper canopy leaves to extreme shade where the PPFD was 10% of full sun with a far-red/red ratio greater than 6. Highly shaded leaves in the high plant density treatment accumulated triacylglycerol (TG) up to 25% of total leaf lipid, a 2.4-fold increase in TG on a chlorophyll basis compared to leaves in the upper canopy. Although total polar lipid content was reduced up to 50% in shaded leaves, shade had little affect on the lipid content or composition of thylakoid membranes. Shade did not affect leaf chlorophyll content. Therefore, the changes in leaf lipid composition were not related to senescence. These findings suggest that conditions of low irradiance and/or a high FR/R ratio cause a shift in carbon metabolism toward the accumulation of TG, a storage lipid. Eighteen-carbon fatty acid desaturation was also affected in highly shaded leaves where a reduction in linolenic acid (18:3) content was accompanied by a proportional increase in oleic (18:1) and linoleic (18:2) acids.     

Induction of CO<Subscript>2</Subscript>-gas exchange and electron transport: comparison of dynamic and steady-state responses in <Emphasis Type="Italic">Fagus sylvatica </Emphasis>leaves

Photosynthetic induction was followed in a juvenile understorey beech (Fagus sylvatica L.) in a shaded habitat which was temporarily exposed to direct sunlight passing through a gap in the canopy. Simultaneous in situ measurements of leaf gas exchange and chlorophyll fluorescence were carried out and a steady-state light response curve of photosynthesis was recorded. The measured dynamic carbon gain was compared to the predicted carbon gain, calculated from the steady-state light response curve and the ambient photon flux density (PPFD) incident during the sunfleck event. Integration over the first 20 min, during which induction took place, resulted in a deviation of 27% if any induction effects are disregarded. The carbon gain was overestimated by 11% when the carbon gain was integrated over the whole measuring period of about 1 h including a 40-min period of full induction. In situ gas exchange measurements under constant saturating light, following dark adaptation, revealed an induction error of 21%. About 20 min was required to reach the final steady-state level of photosynthesis. The data show that the prediction of the carbon gain by steady-state models leads to a distinct overestimation of the CO₂ uptake, irrespective of whether the induction state rises concurrently with the incident radiation or saturating light induces photosynthesis. From chlorophyll fluorescence and absorptance data, rates of linear electron transport (ETR) were calculated. Uninduced leaves exposed to saturating light of constant PPFD show an initial fast down-regulation of ETR, due to excessive light, and a subsequent increase in electron transport attributable to the increasing energy demand during induction of carbon fixation. No difference in the ETR to PPFD relationship between data sets sampled during induction and in the fully induced state was found when both the incident light and the induction state of carbon fixation increased concomitantly. The proportion of electrons fed into alternative pathways besides carbon fixation was higher during induction as compared to the fully induced state. Thus, electrons were used for carbon fixation with a higher efficiency when full induction was reached.     

Leaf construction cost, nutrient concentration, and net CO2 assimilation of native and invasive species in Hawaii

To examine the predictability of leaf physiology and biochemistry from light gradients within canopies, we measured photosynthetic light-response curves, leaf mass per area (LMA) and concentrations of nitrogen, phosphorus and chlorophyll at 15–20 positions within canopies of three conifer species with increasing shade tolerance, ponderosa pine [Pinus ponderosa (Laws.)], Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. Adjacent to each sampling position, we continuously monitored photosynthetically active photon flux density (PPFD) over a 5-week period using quantum sensors. From these measurements we calculated FPAR: integrated PPFD at each sampling point as a fraction of full sun. From the shadiest to the brightest canopy positions, LMA increased by about 50% in ponderosa pine and 100% in western hemlock; Douglas fir was intermediate. Canopy-average LMA increased with decreasing shade tolerance. Most foliage properties showed more variability within and between canopies when expressed on a leaf area basis than on a leaf mass basis, although the reverse was true for chlorophyll. Where foliage biochemistry or physiology was correlated with FPAR, the relationships were non-linear, tending to reach a plateau at about 50% of full sunlight. Slopes of response functions relating physiology and biochemistry to ln(FPAR) were not significantly different among species except for the light compensation point, which did not vary in response to light in ponderosa pine, but did in the other two species. We used the physiological measurements for Douglas fir in a model to simulate canopy photosynthetic potential (daily net carbon gain limited only by PPFD) and tested the hypothesis that allocation of carbon and nitrogen is optimized relative to PPFD gradients. Simulated photosynthetic potential for the whole canopy was slightly higher (<10%) using the measured allocation of C and N within the canopy compared with no stratification (i.e., all foliage identical). However, there was no evidence that the actual allocation pattern was optimized on the basis of PPFD gradients alone; simulated net carbon assimilation increased still further when even more N and C were allocated to high-light environments at the canopy top. Received: 12 August 1998 / Accepted: 25 March 1999     

Photosynthetic performance of the aquatic macrophyte <Emphasis Type="Italic">Althenia orientalis</Emphasis> to solar radiation along its vertical stems

We have studied the plasticity of the photosynthetic apparatus in the endangered aquatic macrophyte Althenia orientalis to the gradient of light availability within its meadow canopy. We determined diurnal change in situ irradiance, light quality, in vivo chlorophyll a fluorescence, ex situ oxygen evolution rates, respiration rate and pigment concentration. The levels of photosynthetic photon flux density (PFD) and ultraviolet radiation (UVR) and the red/far-red ratio decreased with depth within the canopies of A. orientalis. Apical leaves had a greater decrease of the maximal quantum yield (F _v/F _m) in the morning and a faster recovery rate in the afternoon than those in the basal ones. The relative electron transport rate (ETRr) was not saturated at any time of the day, even in the apical leaves that received the highest light. The maximum light-saturated rate of gross photosynthesis (GP_max) took place in apical leaves around noon. The chlorophyll a/b ratio values were higher, and the chlorophyll/carotenoid ratio values lower, in apical leaves than basal ones. The highest concentrations in total carotenoids were reached in the apical leaves around noon. A. orientalis has a high capacity to acclimatize to the changes in the light environment, both in quality and quantity, presenting sun and shade leaves in the same stem through the vertical gradient in the canopy.     

Effets du rapport RC:RS sur les jeunes feuilles de trèfle blanc. Assimilation nette de CO2, activité photochimique et morphologie foliaire

Young leaves of white clover are subjected to low irradiance and low red to far-red (R:FR) ratio within canopies. The objectives were to investigate the consequences of low R:FR ratio on morphology, net CO₂ assimilation and photochemical activity of leaves developed under simulated light environment of canopy. We used far-red (FR) light emitting diodes to modify the R:FR ratio only at the developing leaf under a low irradiance. Net CO₂ assimilation rate, stomatal conductance and leaf morphology were not affected by low R:FR ratio. FR exposure slightly reduced the photochemical quantum yield of PSII but there were no consequences on electron flow through photosystem II. The carbon fixation by the leaf was therefore not modified by light quality. However, low R:FR ratio decreased the leaf chlorophyll content by 21 %. Those effects were only attributed to just unfolded leaves as they were not persistent in mature leaves and there were no consequences on plant biomass accumulation.     

Light and VPD gradients drive foliar nitrogen partitioning and photosynthesis in the canopy of European beech and silver fir

While foliar photosynthetic relationships with light, nitrogen, and water availability have been well described, environmental factors driving vertical gradients of foliar traits within forest canopies are still not well understood. We, therefore, examined how light availability and vapour pressure deficit (VPD) co-determine vertical gradients (between 12 and 42 m and in the understorey) of foliar photosynthetic capacity (Amax), 13C fractionation (∆), specific leaf area (SLA), chlorophyll (Chl), and nitrogen (N) concentrations in canopies of Fagus sylvatica and Abies alba growing in a mixed forest in Switzerland in spring and summer 2017. Both species showed lower Chl/N and lower SLA with higher light availability and VPD at the top canopy. Despite these biochemical and morphological acclimations, Amax during summer remained relatively constant and the photosynthetic N-use efficiency (PNUE) decreased with higher light availability for both species, suggesting suboptimal N allocation within the canopy. ∆ of both species were lower at the canopy top compared to the bottom, indicating high water-use efficiency (WUE). VPD gradients strongly co-determined the vertical distribution of Chl, N, and PNUE in F. sylvatica, suggesting stomatal limitation of photosynthesis in the top canopy, whereas these traits were only related to light availability in A. alba. Lower PNUE in F. sylvatica with higher WUE clearly indicated a trade-off in water vs. N use, limiting foliar acclimation to high light and VPD at the top canopy. Species-specific trade-offs in foliar acclimation to environmental canopy gradients may thus be considered for scaling photosynthesis from leaf to canopy to landscape levels.     

Light-associated nitrogen distribution profile in flowering canopies of sunflower (Helianthus annuus L.) altered during grain growth

In vegetative canopies of many species, the vertical gradient of lamina nitrogen concentration (NW) parallels the profile of light distribution in such a way that the actual nitrogen partitioning approaches the optimum pattern for canopy photosynthesis. This paper evaluates the hypothesis that a strong sink for nitrogen, viz. growing grain, affects the pattern of lamina nitrogen distribution usually described for vegetative canopies. The light and NW profiles of sunflower (Helianthus annuus L.) crops were characterised from anthesis to physiological maturity. The factorial combination of two plant populations (2.4 and 4.8 plants m^–2) and two levels of nitrogen supply (0 and 5 g N m^–2) were the sources of variation for NW and light profiles. Before the onset of nitrogen accumulation in grain, the pattern of NW was similar to that described for other species and it was related to the distribution of light in the canopy. Important changes in the profile of NW occurred during grain filling that were unrelated to the light regime. Nitrogen was mobilised from leaves in all positions in the canopy and the rate of NW change was greater in leaves closer to the grain, which were also the leaves where nitrogen was more concentrated. It is concluded that the physiological mechanisms involved in determining the distribution of leaf nitrogen in vegetative canopies do not apply to sunflower during grain filling.     

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     

Photomodulation of axis extension in sparse canopies : role of the stem in the perception of light-quality signals of stand density

A fiber optic probe inserted into plant tissues was used to investigate the effects of canopy density on the light environment in different organs. The red:far-red ratio inside the stem of Datura ferox L. seedlings and the estimated phytochrome photoequilibrium were strongly reduced by the presence of neighbors forming canopies too sparse to cause any mutual shading at the level of the leaves. In such canopies, changes in plant density had little effects on the light regime inside the leaves of the succulent Aeonium haworthii (S.D.) Webb et Berth., particularly when the lamina was kept nearly normal to the direct rays of the sun. In field experiments using D. ferox and Sinapis alba L. seedlings, the elongation of the internodes responded to various types of localized light-quality treatments that simulated different plant densities in sparse canopies. The responses were quantitatively similar to those elicited by changes in plant density. The evidence supports the hypothesis that, in stands formed by plants of similar size, the red:far-red ratio of the light that impinges laterally on the stems is among the earliest environmental cues that allow plants to detect local canopy density and adjust axis extension accordingly.     

Density-dependent regulation of ramet recruitment by the red:far-red ratio of solar radiation: a field evaluation with the bunchgrass Schizachyrium scoparium

Depressions in the red to far-red ratio (R:FR) of solar radiation arising from the selective absorption of R (600–700 nm) and scattering of FR (700–800 nm) by chlorophyll within plant canopies may function as an environmental signal directly regulating axillary bud growth and subsequent ramet recruitment in clonal plants. We tested this hypothesis in the field within a single cohort of parental ramets in established clones of the perennial bunchgrass, Schizachyrium scoparium. The R:FR was modified near leaf sheaths and axillary buds at the bases of individual ramets throughout the photoperiod without increasing photosynthetic photon flux density (PPFD) by either (1) supplementing R beneath canopies to raise the naturally low R:FR or (2) supplementing FR beneath partially defoliated canopies to suppress the natural R:FR increase following defoliation. Treatment responses were assessed by simultaneously monitoring ramet recruitment, PPFD and the R:FR beneath individual clone canopies at biweekly intervals over a 12-week period. Neither supplemental R nor FR influenced the rate or magnitude of ramet recruitment despite the occurrence of ramet recruitment in all experimental clones. In contrast, defoliation with or without supplemental FR beneath clone canopies reduced ramet recruitment 88% by the end of the experiment. The hypothesis stating that the R:FR signal directly regulates ramet recruitment is further weakened by evidence demonstrating that (1) the low R:FR-induced suppression of ramet recruitment is only one component of several architectural modifications exhibited by ramets in response to the R:FR signal (2) immature leaf blades, rather than leaf sheaths or buds, function as sites of R:FR perception on individual ramets, and (3) increases in the R:FR at clone bases following partial canopy removal are relatively transient and do not override the associated constraints on ramet recruitment resulting from defoliation. A depressed R:FR is probably of greater ecological significance as a signal of competition for light in vegetation canopies than as a density-dependent signal which directly regulates bud growth and ramet recruitment.     

Relationships between leaf nitrogen and limitations of photosynthesis in canopies of Solidago altissima

Vertical distribution patterns of light, leaf nitrogen, and leaf gas exchange through canopies of the clonal perennial Solidago altissima were studied in response to mowing and fertilizer application in a field experiment. Consistent with the distribution of light, average leaf nitrogen content followed a `smooth' exponential decline along the fertilized stands both in control and mown plots. The nitrogen profile along the unfertilized stands in mown plots, however, was `disrupted' by high-nitrogen leaves at the top of shorter ramets that only reached intermediate strata of the canopies. Hence, in these stands leaf nitrogen was significantly increased in short ramets compared with tall ramets for a given light environment, suggesting suboptimal stand structure but not necessarily suboptimal single-ramet architecture. However, at least under the climatic conditions observed during measurements, such disrupture had no substantial effect on stand productivity: model calculations showed that vertical distribution patterns of leaf nitrogen along ramets only marginally influenced the photosynthetic performance of ramets and stands. This is explained by the observed photosynthesis-nitrogen relationship: the rate of photosynthesis per unit amount of leaf nitrogen did not increase with leaf nitrogen content even under saturating light levels indicating that leaf photosynthesis was not nitrogen limited during the measurement periods. Nevertheless, our study indicates that consideration of how architecture(s) of adjacent individual plants interact could be essential for a better understanding of the trade-offs between individual and canopy characteristics for maximizing carbon gain. Such trade-offs may end up in a suboptimal canopy structure, which could not be predicted and understood by classical canopy optimization models.     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

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.     

Under slow flow conditions,daily rates of canopy photosynthesis of marine macrophytes were insensitive to model choice and flow‐rate

Water motion drives the flux of suspended and dissolved material (e.g., nutrients, gametes, and dissolved oxygen) to and from macrophyte canopies, and is one of the most important mechanisms that can regulate the growth, survival, and persistence of marine macrophytes populations. At small spatial scales (e.g., lamina or leaves and individuals), increasing flow‐rates have been demonstrated to enhance physiological processes, especially photosynthesis rates, and we expected a similar response at the canopy scale. We conducted seven experiments over 25 days using a pair of open‐air flow‐chambers under natural light, temperature, and seawater conditions. In the four marine macrophyte (Sargassum piluliferum, S. siliquastrum, S. thunbergii, and Zostera marina) canopies examined, an increase in flow‐rate did not enhance photosynthesis rates. The odds that daily gross photosynthesis rates increase with a decrease in flow‐rates was 1.77 to 1. We also examined if two non‐linear equations and one linear equation, often used to describe the relationship between photosynthesis to photosynthetic photon flux density (PPFD), biased estimates of the daily rates of photosynthesis and respiration. It was revealed that the functional form of the equation strongly influenced photosynthesis and respiration rate estimates at short time scales (i.e., minutes), however, daily rates were insensitive to the type of equation used to model the relationship between photosynthesis and PPFD. We suggest that the predominance of photosynthesis rates occurring in under‐saturating PPFD conditions (> 40 % of daylight hours) may be one of the reasons for this insensitivity.     

Photomorphogenesis, Photosynthesis, and Early Growth of Primary Leaves of Phaseolus vulgaris

The response of leaf tissue to white, blue, red, and far-redlight has been examined. Leaves on plants grown in darknessshow increased cell number, cell volume, and area when exposedto long periods (up to 48 h) of low-intensity red, blue, orfar-red radiation. This is believed to be a photomorphogenicresponse which does not involve photosynthesis. Leaves fromplants exposed to white light during germination do not usuallyrespond to red, blue, or far-red light. An exception to thiswas found for leaf discs which showed a larger increase in areathan the dark controls following exposure to far-red light for24 h. Leaf tissue from light-grown plants responds to high-intensitywhite light, probably through photosynthesis. Discs cut fromdark-grown plants and cultured in white light grow equally wellin air and CO₂-free conditions. Application of the photosyntheticinhibitor DCMU reduces growth and chlorophyll formation, however. It is concluded that light, perhaps acting through the phytochromemechanism, has initially a number of morphogenic effects includinginitiation of development of the photosynthetic apparatus. Theresponses to photomorphogenically active radiation do not persistand light effects through photosynthesis are rapidly initiatedand dominate the later stages of leaf growth.