A new paradigm in leaf-level photosynthesis: direct and diffuse lights are not equal

Global-change scenarios suggest a trend of increasing diffuse light due to expected increases in cloud cover. Canopy-level measurements of plant-community photosynthesis under diffuse light show increased productivity attributed to more uniform distribution of light within the forest canopy, yet the effect of the directional quality of light at the leaf level is unknown. Here we show that leaf-level photosynthesis in sun leaves of both C(3) and C(4) plants can be 10-15% higher under direct light compared to equivalent absorbed irradiances of diffuse light. High-light-grown leaves showed significant photosynthetic enhancement in direct light, while shade-adapted leaves showed no preference for direct or diffuse light at any irradiance. High-light-grown leaves with multiple palisade layers may be adapted to better utilize direct than diffuse light, while shade leaf structure does not appear to discriminate light based on its directionality. Based upon our measurements, it appears that leaf-level and canopy-level photosynthetic processes react differently to the directionality of light, and previously observed increases in canopy-level photosynthesis occur even though leaf-level photosynthesis decreases under diffuse light.     

Photosynthetic acclimation to simultaneous and interacting environmental stresses along natural light gradients: optimality and constraints

There is a strong natural light gradient from the top to the bottom in plant canopies and along gap-understorey continua. Leaf structure and photosynthetic capacities change close to proportionally along these gradients, leading to maximisation of whole canopy photosynthesis. However, other environmental factors also vary within the light gradients in a correlative manner. Specifically, the leaves exposed to higher irradiance suffer from more severe heat, water, and photoinhibition stresses. Research in tree canopies and across gap-understorey gradients demonstrates that plants have a large potential to acclimate to interacting environmental limitations. The optimum temperature for photosynthetic electron transport increases with increasing growth irradiance in the canopy, improving the resistance of photosynthetic apparatus to heat stress. Stomatal constraints on photosynthesis are also larger at higher irradiance because the leaves at greater evaporative demands regulate water use more efficiently. Furthermore, upper canopy leaves are more rigid and have lower leaf osmotic potentials to improve water extraction from drying soil. The current review highlights that such an array of complex interactions significantly modifies the potential and realized whole canopy photosynthetic productivity, but also that the interactive effects cannot be simply predicted as composites of additive partial environmental stresses. We hypothesize that plant photosynthetic capacities deviate from the theoretical optimum values because of the interacting stresses in plant canopies and evolutionary trade-offs between leaf- and canopy-level plastic adjustments in light capture and use.     

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

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

Consumer Depletion Alters Seagrass Resistance to an Invasive Macroalga

Few field studies have investigated how changes at one trophic level can affect the invasibility of other trophic levels. We examined the hypothesis that the spread of an introduced alga in disturbed seagrass beds with degraded canopies depends on the depletion of large consumers. We mimicked the degradation of seagrass canopies by clipping shoot density and reducing leaf length, simulating natural and anthropogenic stressors such as fish overgrazing and water quality. Caulerpa racemosa was transplanted into each plot and large consumers were excluded from half of them using cages. Potential cage artifacts were assessed by measuring irradiance, scouring by leaf movement, water flow, and sedimentation. Algal invasion of the seagrass bed differed based on the size of consumers. The alga had higher cover and size under the cages, where the seagrass was characterized by reduced shoot density and canopy height. Furthermore, canopy height had a significant effect depending on canopy density. The alteration of seagrass canopies increased the spread of C. racemosa only when large consumers were absent. Our results suggest that protecting declining habitats and/or restoring fish populations will limit the expansion of C. racemosa. Because MPAs also enhance the abundance and size of fish consuming seagrass they can indirectly promote algal invasion. The effects of MPAs on invasive species are context dependent and require balancing opposing forces, such as the conservation of seagrass canopy structure and the protection of fish grazing the seagrass.     

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.     

温室栽培西瓜座果期冠层光合作用日变化研究

在兰州春季塑料日光温室条件下研究了‘宝冠’和‘新金兰’两个品种西瓜(Citrullus lantus(Thunb．) Mansf．)冠层光合作用日变化特性。结果表明,日光温室两个西瓜品种间冠层光照分布和单叶光合作用日变化模式存在明显的差异,冠层光照条件影响各叶层的光合强度及其日变化态势。两品种冠层整体光合速率日动态呈单峰型,‘宝冠’的光合生产力高于‘新金兰’。‘宝冠’西瓜单叶光合速率较高,高效光合时间较长,且表现出午休特征。‘新金兰’西瓜单叶光合速率低,高效光合维持时间较短,未表现出明显午休特征。‘宝冠’光合作用的瞬时水分效率 (WUE)高于‘新金兰’,两品种上午的WUE高于下午,上层叶的WUE高于下层叶。     

Canopy Light Gradient Perception by Cytokinin

We have recently identified cytokinin as an important xylem-carried signal involved in the photosynthetic acclimation of plants to light gradients in dense canopies. Lower leaves become shaded in a dense canopy and consequently have reduced transpiration rates. our measurements have shown that this results in a reduced delivery of cytokinins carried in the transpiration stream to shaded leaves, as compared to light-exposed leaves. Cytokinins are involved in the regulation of photosynthetic acclimation to the light gradient by stimulating the expression of photosynthetic enzymes in light-exposed leaves. In shaded leaves, the low delivery rate of cytokinin leads to reduced photosynthetic capacity and ultimately senescence. We show evidence for this role of cytokinin, as part of a complex of signaling pathways where other regulatory mechanisms are also involved. A model is presented depicting the regulation of photosynthetic acclimation by cytokinin delivery to leaves dependent on the irradiance they receive.Key Words: canopy light gradient, transpiration, photosynthetic acclimation, cytokinin, nitrate, systemic signaling     

Altered physiological function,not structure,drives increased radiation-use efficiency of soybean grown at elevated CO<Subscript>2</Subscript>

Previous studies of elevated carbon dioxide concentration ([CO₂]) on crop canopies have found that radiation-use efficiency is increased more than radiation-interception efficiency. It is assumed that increased radiation-use efficiency is due to changes in leaf-level physiology; however, canopy structure can affect radiation-use efficiency if leaves are displayed in a manner that optimizes their physiological capacity, even though the canopy intercepts the same amount of light. In order to determine the contributions of physiology and canopy structure to radiation-use and radiation-interception efficiency, this study relates leaf-level physiology and leaf display to photosynthetic rate of the outer canopy. We used a new imaging approach that delivers three-dimensional maps of the outer canopy during the growing season. The 3D data were used to model leaf orientation and mean photosynthetic electron transport of the outer canopy to show that leaf orientation changes did not contribute to increased radiation-use; i.e. leaves of the outer canopy showed similar diurnal leaf movements and leaf orientation in both treatments. Elevated [CO₂] resulted in an increased maximum electron transport rate (ETR_max) of light reactions of photosynthesis. Modeling of canopy light interception showed that stimulated leaf-level electron transport at elevated [CO₂], and not alterations in leaf orientation, was associated with stimulated radiation-use efficiency and biomass production in elevated [CO₂]. This study provides proof of concept of methodology to quantify structure–function relationships in combination, allowing a quantitative estimate of the contribution of both effects to canopy energy conversion under elevated [CO₂].     

The responses of light interception,photosynthesis and fruit yield of cucumber to LED‐lighting within the canopy

Mathematical models of light attenuation and canopy photosynthesis suggest that crop photosynthesis increases by more uniform vertical irradiance within crops. This would result when a larger proportion of total irradiance is applied within canopies (interlighting) instead of from above (top lighting). These irradiance profiles can be generated by Light Emitting Diodes (LEDs). We investigated the effects of interlighting with LEDs on light interception, on vertical gradients of leaf photosynthetic characteristics and on crop production and development of a greenhouse‐grown Cucumis sativus‘Samona’ crop and analysed the interaction between them. Plants were grown in a greenhouse under low natural irradiance (winter) with supplemental irradiance of 221 µmol photosynthetic photon flux m^?2 s^?1 (20 h per day). In the interlighting treatment, LEDs (80% Red, 20% Blue) supplied 38% of the supplemental irradiance within the canopy with 62% as top lighting by High‐Pressure Sodium (HPS)‐lamps. The control was 100% top lighting (HPS lamps). We measured horizontal and vertical light extinction as well as leaf photosynthetic characteristics at different leaf layers, and determined total plant production. Leaf mass per area and dry mass allocation to leaves were significantly greater but leaf appearance rate and plant length were smaller in the interlighting treatment. Although leaf photosynthetic characteristics were significantly increased in the lower leaf layers, interlighting did not increase total biomass or fruit production, partly because of a significantly reduced vertical and horizontal light interception caused by extreme leaf curling, likely because of the LED‐light spectrum used, and partly because of the relatively low irradiances from above.     

Comparative ecophysiology of leaf and canopy photosynthesis

Leaves and herbaceous leaf canopies photosynthesize efficiently although the distribution of light, the ultimate resource of photosynthesis, is very biased in these systems. As has been suggested in theoretical studies, if a photosynthetic system is organized such that every photosynthetic apparatus photosynthesizes in concert, the system as a whole has the sharpest light response curve and is most adaptive. This condition can be approached by (i) homogenization of the light environment and (ii) acclimation of the photosynthetic properties of leaves or chloroplasts to their local light environments. This review examines these two factors in the herbaceous leaf canopy and in the leaf. Changes in the inclination of leaves in the canopy and differentiation of mesophyll into palisade and spongy tissue contribute to the moderation of the light gradient. Leaf and chloroplast movements in the upper parts of these systems under high irradiances also moderate light gradients. Moreover, acclimation of leaves and chloroplasts to the local light environment is substantial. These factors increase the efficiency of photosynthesis considerably. However, the systems appear to be less efficient than the theoretical optimum. When the systems are optically dense, the light gradients may be too great for leaves or chloroplasts to acclimate. The loss of photosynthetic production attributed to the imperfect adjustment of photosynthetic apparatus to the local light environment is most apparent when the photosynthesis of the system is in the transition between the light-limited and light-saturated phases. Although acclimation of the photosynthetic apparatus and moderation of light gradients are imperfect, these markedly raise the efficiency of photosynthesis. Thus more mechanistic studies on these adaptive attributes are needed. The causes and consequences of imperfect adjustment should also be investigated.     

Photosynthetic acclimation in relation to nitrogen allocation in cucumber leaves in response to changes in irradiance

Leaves deep in canopies can suddenly be exposed to increased irradiances following e.g. gap formation in forests or pruning in crops. Studies on the acclimation of photosynthesis to increased irradiance have mainly focused on the changes in photosynthetic capacity (A_max), although actual irradiance often remains below saturating level. We investigated the effect of changes in irradiance on the photosynthesis irradiance response and on nitrogen allocation in fully grown leaves of Cucumis sativus. Leaves that fully developed under low (50 µmol m^?2 s^?1) or moderate (200 µmol m^?2 s^?1) irradiance were subsequently exposed to, respectively, moderate (LM‐leaves) or low (ML‐leaves) irradiance or kept at constant irradiance level (LL‐ and MM‐leaves). Acclimation of photosynthesis occurred within 7 days with final A_max highest in MM‐leaves, lowest in LL‐leaves and intermediate in ML‐ and LM‐leaves, whereas full acclimation of thylakoid processes underlying photosystem II (PSII) efficiency and non‐photochemical quenching occurred in ML‐ and LM‐leaves. Dark respiration correlated with irradiance level, but not with A_max. Light‐limited quantum efficiency was similar in all leaves. The increase in photosynthesis at moderate irradiance in LM‐leaves was primarily driven by nitrogen import, and nitrogen remained allocated in a similar ratio to Rubisco and bioenergetics, while allocation to light harvesting relatively decreased. A contrary response of nitrogen was associated with the decrease in photosynthesis in ML‐leaves. Net assimilation of LM‐leaves under moderate irradiance remained lower than in MM‐leaves, revealing the importance of photosynthetic acclimation during the leaf developmental phase for crop productivity in scenarios with realistic, moderate fluctuations in irradiance that leaves can be exposed to.     

Instantaneous canopy photosynthesis: analytical expressions for sun and shade leaves based on exponential light decay down the canopy and an acclimated non-rectangular hyperbola for leaf photosynthesis

Analytical expressions for the contributions of sun and shade leaves to instantaneous canopy photosynthesis are derived. The analysis is based on four assumptions. First, that the canopy is closed in the sense that it is horizontally uniform. Secondly, that there is an exponential profile of light down the canopy with the same decay constant for light from different parts of the sky. Thirdly, that the leaf photosynthetic response to incident irradiance can be described by a three-parameter non-rectangular hyperbola (NRH). And lastly, that light acclimation at the leaf level occurs in only one parameter of the NRH, that describing the light-saturated photosynthetic rate, which is assumed to be proportional to the local averaged leaf irradiance. These assumptions have been extensively researched empirically and theoretically and their limitations are quite well understood. They have been widely used when appropriate. Combining these four assumptions permits the derivation of algebraic expressions for instantaneous canopy photosynthesis which are computationally efficient because they avoid the necessity for numerical integration down the canopy. These are valuable for modelling plant and crop ecosystems, for which canopy photosynthesis is the primary driver. Ignoring the sun/shade dichotomy can result in overestimates of canopy photosynthesis of up to 20 %, but using a rectangular hyperbola instead of a non-rectangular hyperbola to estimate canopy photosynthesis taking account of sun and shade leaves can lead to a similarly sized underestimate.     

Negative feedback regulation is responsible for the non-linear modulation of photosynthetic activity in plants and cyanobacteria exposed to a dynamic light environment

Photosynthetic organisms exposed to a dynamic light environment exhibit complex transients of photosynthetic activities that are strongly dependent on the temporal pattern of the incident irradiance. In a harmonically modulated light of intensity I≈const.+sin(ωt), chlorophyll fluorescence response consists of a steady-state component, a component modulated with the angular frequency of the irradiance ω and several upper harmonic components (2ω, 3ω and higher). Our earlier reverse engineering analysis suggests that the non-linear response can be caused by a negative feedback regulation of photosynthesis. Here, we present experimental evidence that the negative feedback regulation of the energetic coupling between phycobilisome and Photosystem II (PSII) in the cyanobacterium Synechocystis sp. PCC6803 indeed results in the appearance of upper harmonic modes in the chlorophyll fluorescence emission. Dynamic changes in the coupling of the phycobilisome to PSII are not accompanied by corresponding antiparallel changes in the Photosystem I (PSI) excitation, suggesting a regulation limited to PSII. Strong upper harmonic modes were also found in the kinetics of the non-photochemical quenching (NPQ) of chlorophyll fluorescence, of the P700 redox state and of the CO₂ assimilation in tobacco (Nicotiana tabaccum) exposed to harmonically modulated light. They are ascribed to negative feedback regulation of the reactions of the Calvin-Benson cycle limiting the photosynthetic electron transport. We propose that the observed non-linear response of photosynthesis may also be relevant in a natural light environment that is modulated, e.g., by ocean waves, moving canopy or by varying cloud cover. Under controlled laboratory conditions, the non-linear photosynthetic response provides a new insight into dynamics of the regulatory processes.     

Sunspots drive seagrasses

Seagrasses are underwater flowering plants that form an important marine habitat worldwide. They respond to watershed and climate influences and are considered a good indicator of environmental health. Investigations into seagrass photosynthesis, response to high irradiance and UV light are providing new insights into controls on plant production. We show that the size and density of the intertidal seagrass directly relates to sunspot activity. The influence of sunspot activity has been largely overlooked in plants and never examined in seagrasses. The correlation between seagrass canopy height and the number of sunspots was highly significant, with height decreasing steadily as sunspots increase and recover only after a substantial decline in sunspot activity. The density of seagrass shoots correlated positively with sunspots until, at high sunspot numbers (>110 per month), plant density dropped and then rebounded. High sunspot activity inhibits the production and alters the meadow structure of tropical intertidal seagrass habitats.     

Dynamics of Vertical Leaf Nitrogen Distribution in a Vegetative Wheat Canopy. Impact on Canopy Photosynthesis

The development of vertical canopy gradients of leaf N has beenregarded as an adaptation to the light gradient that helps tomaximize canopy photosynthesis. In this study we report thedynamics of vertical leaf N distribution during vegetative growthof wheat in response to changes in N availability and sowingdensity. The question of to what extent the observed verticalleaf N distribution maximized canopy photosynthesis was addressedwith a leaf layer model of canopy photosynthesis that integratesN-dependent leaf photosynthesis according to the canopy lightand leaf N distribution. Plants were grown hydroponically attwo amounts of N, supplied in proportion to calculated growthrates. Photosynthesis at light saturation correlated with leafN. The vertical leaf N distribution was associated with thegradient of absorbed light. The leaf N profile changed duringcrop development and was responsive to N availability. At highN supply, the leaf N profiles were constant during crop development.At low N supply, the leaf N profiles fluctuated between moreuniform and steep distributions. These changes were associatedwith reduced leaf area expansion and increasing N remobilizationfrom lower leaf layers. The distribution of leaf N with respectto the gradient of absorbed irradiance was close to the theoreticaloptimum maximizing canopy photosynthesis. Sensitivity analysisof the photosynthesis model suggested that plants maintain anoptimal vertical leaf N distribution by balancing the capacityfor photosynthesis at high and low light. Copyright 2000 Annalsof Botany Company Canopy photosynthesis, leaf nitrogen distribution, nitrogen, Triticum aestivum L, wheat     

Leaf photosynthetic light response: a mechanistic model for scaling photosynthesis to leaves and canopies

1. The response of photosynthesis to radiation is an often-studied but poorly understood process, represented empirically in most photosynthesis models. However, in scaling photosynthesis from leaf to canopy, predictions of canopy photosynthesis are very sensitive to parameters describing the response of leaves to Photosynthetic Photon Flux Density (PPFD).
2. In this study, a mechanistic, yet still simple, approach is presented that models the degree of light saturation in leaves explicitly, assuming a heterogeneous environment of PPFD and chlorophyll.
3. Possible mechanisms determining the ratio of chlorophyll to nitrogen are considered, including a direct dependence on PPFD, a mechanism involving the red/far-red ratio of light in the canopy, and an approach based upon maximizing photosynthesis.
4. Comparison of model predictions with two data sets of light, nitrogen and chlorophyll from canopies of Populus and Corylus suggests that the red/far-red mechanism is the most realistic. The data also show that the trees studied do not always optimize their nitrogen partitioning to maximize photosynthetic yield.
5. We then apply the model to the data sets, to predict the shape of light response curves of leaves within canopies and assess the applicability of simple scaling schemes, in which full acclimation of photosynthesis to PPFD justifies the use of big-leaf models. We conclude that, at least for the data used, basic assumptions of such schemes do not hold.     

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     

CHARACTERIZING AND QUANTIFYING PHOTOINHIBITION IN INTERTIDAL MICROPHYTOBENTHOS1

This study characterizes the short‐term influence of the sustained saturating irradiance encountered by the microphytobenthos inhabiting intertidal mudflats. The kinetics of photoinhibition in epipelic microalgae from intertidal mudflats were investigated in the laboratory. Previously isolated benthic microalgae were exposed to a saturating photon flux density (PFD) for periods ranging from 0 to 180 min; every 30 min, a photosynthesis‐irradiance curve was established to quantify the effect of the saturating PFD on both parameters α^B, the photosynthetic efficiency, and P_m^B, the photosynthetic capacity. The α^B decreased from the beginning of light exposure until the end, whereas P_m^B first slightly increased and then diminished from 90 min exposure onward. It turned out that epipelic microphytobenthos undergoes photoinhibition after about 90 min of saturating PFD. The possible ecological consequences of these ecophysiological results are discussed.     

Light-Induced Adaptive Responses under Greenhouse and Controlled Conditions in the Fern Pteris cretica var. ouvrardii

The photosynthetic capabilities of the fern Pteris cretica var. ouvrardii were analysed by means of the light response curves of CO₂ exchange. In control growth conditions (greenhouse, low-light: 20–32 W m^?2); photosynthesis was shown to be saturated for low irradiance (20–25 W m^?2); the saturating photosynthetic rate, very low as compared to higher plants, was due to an extremely high intracellular resistance. When irradiance during the photosynthesis measurement was higher than 60–80 W m^?2, a constant decline of net CO₂ exchange as a function of time was observed. When irradiance during growth was enhanced, whether in greenhouse (20–250 W m^?2) or controlled (62 W m^?2) conditions, the first fronds that had developed in the new condition from the crosier stage exhibited decreased net maximal photosynthesis and a decreased efficiency in low light, but saturating irradiance was unmodified. However, the fronds whose entire differentiation (from meristem) occurred under these moderate irradiances (plants defoliated of all fronds and crosiers at the time of transfer), possessed more efficient photosynthetic characteristics than control plants. Pteris is able to grow under extreme shade conditions (4–8 W m^?2); light saturating photosynthesis and efficiency are higher under extreme shade than under control conditions. These adaptive characteristics indicate that Pteris is a well-adapted shade species.     

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.