The Heat and Mass Transfer Analysis of a Leaf

Understanding the heat and mass transfer processes of plant leaves is essential for plant bionic engineering. A general thermophysical model was established for a plant leaf with particular emphasis on the transpiration process. The model was verified by the field measured stomatal resistance and temperature of a camphor leaf. A dynamical simulation revealed that diurnal transpiration water consumption is dominated by the solar irradiance and the day-average temperature of the leaf is dominated by the ambient air temperature; transpiration plays an important role in the cooling of the leaf, in average it could dissipate around 32.9% of the total solar energy absorbed by the leaf in summer. To imitate the thermal infared characteristic of the real leaf, the up surface of the bionic leaf must have emissivity and solar absorptivity close to those of a real leaf and its shape and surface roughness must be similar to those of the real leaf. The key point is that the bionic leaf must be able to evaporate water to simulate the transpiration of a plant leaf, appropriate adsorbent can be used to realize this function.     

夏玉米叶片水分变化与光合作用和土壤水分的关系

叶片是光合作用的重要器官,其含水量的变化必将影响光合作用,但关于叶片水分变化对光合作用的影响报道较少。以华北夏玉米为研究对象,利用三叶期不同水分梯度的持续干旱模拟试验资料,分析夏玉米叶片水分变化及其与叶片净光合速率和土壤水分的关系。结果表明:夏玉米叶片净光合速率对叶片水分变化的响应显著且呈二次曲线关系,叶片含水量约为70.30%时,叶片净光合速率为零;叶片含水量与土壤相对湿度呈非直角双曲线关系,叶片最大含水量约为85.14%。研究结果可为准确描述叶片水分变化对光合作用的影响及客观辨识夏玉米干旱的发生发展及监测预警提供参考。     

夏玉米三叶期持续干旱下不同叶位叶片含水量变化及其与光合作用的关系

植物干物质的累积依赖于群体光合速率,而群体光合速率又与单叶的光合能力密切有关。叶片光合作用与其含水量密切相关,目前关于不同叶位叶片含水量对持续干旱的响应及其与光合作用的关系还未见报道。以华北夏玉米郑单958为材料,设置6个不同灌水处理,模拟不同灌溉量下持续干旱对夏玉米不同叶位叶片生理特性的影响,分析夏玉米顶部开始的第一、三、五叶位叶片的水分变化及其与净光合速率的关系。结果表明:夏玉米不同叶位的叶片最大含水量不同,且随干旱进程的推进叶片含水量的变化速率也不同,第一叶的叶片含水量下降速率高于第三、第五叶,第一叶的最大含水量高于第三、五叶,且可进行光合产物积累的叶片含水量下限随叶位的增加而增大。同时,第一叶的叶片含水量与土壤水分呈显著相关,且与净光合速率的相关性也非常强。第一叶可进行光合产物积累的叶片水分下限(净光合速率为零时的叶片含水量)最小,表明其耐旱性最强,对干旱具有指导意义。研究结果可为提高冠层光合作用模拟的准确性及夏玉米干旱发生发展的监测预警提供参考。     

The influence of leaf size and shape on leaf thermal dynamics: does theory hold up under natural conditions?

Laboratory studies on artificial leaves suggest that leaf thermal dynamics are strongly influenced by the two‐dimensional size and shape of leaves and associated boundary layer thickness. Hot environments are therefore said to favour selection for small, narrow or dissected leaves. Empirical evidence from real leaves under field conditions is scant and traditionally based on point measurements that do not capture spatial variation in heat load. We used thermal imagery under field conditions to measure the leaf thermal time constant (τ) in summer and the leaf‐to‐air temperature difference (?T) and temperature range across laminae (T_range) during winter, autumn and summer for 68 Proteaceae species. We investigated the influence of leaf area and margin complexity relative to effective leaf width (w_e), the latter being a more direct indicator of boundary layer thickness. Normalized difference of margin complexity had no or weak effects on thermal dynamics, but w_e strongly predicted τ and ?T, whereas leaf area influenced T_range. Unlike artificial leaves, however, spatial temperature distribution in large leaves appeared to be governed largely by structural variation. Therefore, we agree that small size, specifically w_e, has adaptive value in hot environments but not with the idea that thermal regulation is the primary evolutionary driver of leaf dissection.     

叶脉结构与功能及其对叶片经济谱的影响

叶脉由贯穿于叶肉内部的维管组织及其外围机械组织构成,多样化的脉序及网络结构使叶脉系统发生变异和功能分化。该文综述了叶脉系统结构与功能的最新研究进展。通过聚焦叶脉分级系统的结构与功能及其在叶片经济谱(LES)中的重要性,解释叶脉性状与其它叶片功能性状之间的关系及机制。不同等级叶脉在机械支撑与水分运输方面存在功能分化,其中1–3级粗脉在维持叶片形状和叶表面积以及物理支撑方面发挥重要作用,有利于维持叶片最大受光面积;4级及以上细脉具有水分调节功能,它们与气孔相互协调,影响叶片水分运输、蒸腾散热和光合作用速率。叶片生长过程与叶脉发育的动态变化模式决定叶脉密度,并影响叶脉密度与叶片大小之间的关系:叶面积与粗脉密度呈显著负相关,与粗脉直径呈显著正相关,而与细脉密度无关。与叶脉性状相关的叶片经济谱框架模型预测,叶脉密度较高的叶片寿命短、比叶重较小,叶片最大碳同化速率、代谢速率以及资源获取策略潜力较高。     

Leaf patch clamp pressure probe measurements on olive leaves in a nearly turgorless state

The non‐invasive leaf patch clamp pressure (LPCP) probe measures the attenuated pressure of a leaf patch, P_p, in response to an externally applied magnetic force. P_p is inversely coupled with leaf turgor pressure, P_c, i.e. at high P_c values the P_p values are small and at low P_c values the P_p values are high. This relationship between P_c and P_p could also be verified for 2‐m tall olive trees under laboratory conditions using the cell turgor pressure probe. When the laboratory plants were subjected to severe water stress (P_c dropped below ca. 50 kPa), P_p curves show reverse diurnal changes, i.e. during the light regime (high transpiration) a minimum P_p value, and during darkness a peak P_p value is recorded. This reversal of the P_p curves was completely reversible. Upon watering, the original diurnal P_p changes were re‐established within 2–3 days. Olive trees in the field showed a similar turnover of the shape of the P_p curves upon drought, despite pronounced fluctuations in microclimate. The reversal of the P_p curves is most likely due to accumulation of air in the leaves. This assumption was supported with cross‐sections through leaves subjected to prolonged drought. In contrast to well‐watered leaves, microscopic inspection of leaves exhibiting inverse diurnal P_p curves revealed large air‐filled areas in parenchyma tissue. Significantly larger amounts of air could also be extracted from water‐stressed leaves than from well‐watered leaves using the cell turgor pressure probe. Furthermore, theoretical analysis of the experimental P_p curves shows that the propagation of pressure through the nearly turgorless leaf must be exclusively dictated by air. Equations are derived that provide valuable information about the water status of olive leaves close to zero P_c.     

The Effect of Leaf Age on Gas Exchange and Malate Accumulation in C3-CAM Plant Marrubium Frivaldszkyanum (Lamiaceae)

For the first time the expression of C₃ and CAM in the leaves of different age of Marrubium frivaldszkyanum Boiss, is reported. With increasing leaf age a typical C₃ photosynthesis pattern and high transpiration rate were found. In older leaves a shift to CAM occurred and the 24-h transpiration water loss decreased. A correlation was established between leaf area and accumulation of malate. Water loss at early stages of leaf expansion may be connected with the shift to CAM and the water economy of the whole plant.     

油松叶功能性状分布特征及其控制因素

植物叶功能性状能够直接或间接反应植物对环境的适应策略,这种适应策略会间接影响植物的生长、繁殖和生存。目前已有大量关于植物叶功能性状与环境间关系的研究,但这些研究多使用性状平均值代替物种,忽略了性状的种内变化。油松是我国重要的造林树种,对我国陆地生态系统起着重要的作用,研究其叶功能性状与环境的关系有助于更好的的理解种内功能性状与环境间的关系。于2014年6—9月对辽宁、内蒙、北京、山西、陕西、宁夏、青海等地天然油松进行采样并对其重要的8个叶功能性状的分布特征及主控环境因子进行了研究,结果表明:1)油松各叶性状值存在较大的种内变异系数(4.82%—25.85%),除1年生叶碳含量(LCC)油松各叶功能性状值在不同研究地点间差异显著(P0.05);2)油松叶长(LL)、叶厚(LT)、比叶面积(SLA)、气孔密度(SD)、叶氮含量(LNC)存在较弱的经度格局,LT、SD、LNC存在较弱的纬度格局(0.05R~20.3),水热条件的变化以及较大的局部效应是造成这种格局的可能原因;3)控制油松各叶性状分布的主要环境因子各不相同,其中LL主要受到年平均降水量和海拔高度的影响;LT主要受到年均温和土壤体积含水率的影响;SLA主要受到年平均降水量和土壤氮含量的影响;叶干物质含量(LDMC)主要受到水因子的影响;SD主要受到海拔高度的影响;LNC主要受到海拔和水因子的影响;叶磷含量(LPC)主要受到土壤磷含量的影响。     

The Effect of Leaf Water Potential, Leaf Temperature and Light Intensity on Leaf Diffusion Resistance and the Transpiration of Leaves of Malus sylvestris

The relationship between leaf resistance to water vapour diffusion and each of the factors leaf water potential, light intensity and leaf temperature was determined for leaves on seedling apple trees (Malus sylvestris Mill. cv. Granny Smith) in the laboratory. Leaf cuticular resistance was also determined and transpiration was measured on attached leaves for a range of conditions. Leaf resistance was shown to be independent of water potential until potential fell below — 19 bars after which leaf resistance increased rapidly. Exposure of leaves to CO₂-free air extended the range for which resistance was independent of water potential to — 30 bars. The light requirement for minimum leaf resistance was 10 to 20 W m^?2 and at light intensities exceeding these, leaf resistance was unaffected by light intensity. Optimum leaf temperature for minimum diffusion resistance was 23 ± 2°C. The rate of change measured in leaf resistance in leaves given a sudden change in leaf temperature increased as the magnitude of the temperature change increased. For a sudden change of 1°C in leaf temperature, diffusion resistance changed at a rate of 0.01 s cm^?1 min^?1 whilst for a 9°C leaf temperature change, diffusion resistance changed at a rate of 0.1 s cm^?1 min^?1. Cuticular resistance of these leaves was 125 s cm^?1 which is very high compared with resistances for open stomata of 1.5 to 4 s cm^?1 and 30 to 35 s cm^?1 for stomata closed in the dark. Transpiration was measured in attached apple leaves enclosed in a leaf chamber and exposed to a range of conditions of leaf temperature and ambient water vapour density. Peak transpiration of approximately 5 × 10^?6 g cm^?2 s^?1 occurred at a vapour density gradient from the leaf to the air of 12 to 14 g m^?3 after which transpiration declined due presumably to increased stomatal resistance. Leaves in CO₂-free air attained a peak transpiration of 11 × 10^?6 g cm^?2 s^?1 due to lower values of leaf resistance in CO₂ free air. Transpiration then declined in these leaves due to development of an internal leaf resistance (of up to 2 s cm^?1). The internal resistance was masked in leaves at normal CO₂ concentrations by the increase in stomatal resistance.     

Changes in Stem and Leaf Hydraulics Preceding Leaf Shedding in Castanea Sativa L.

This paper describes changes in leaf water status and in stem, petiole and leaf blade hydraulics preceding leaf senescence and shedding in Castanea sativa L. (chestnut). Measurements of maximum diurnal leaf conductance to water vapour (g_L), minimum water potential (_L), hydraulic conductance per unit leaf surface area of stems (K_SL), petioles (K_PL) and leaf blades (K_LL) and number of functional conduits and inside diameter distribution were performed in June, September and October 1999. In September, still green leaves had undergone some dehydration as indicated by decreased g_L (by 75 %) and _L with respect to June. In the same time, K_SL decreased by 88 %, while K_PL and K_LL decreased by 50 % and 20 % of the conduits of stems and 10 % of the petioles (all belonging to the widest diameter range) were no longer functioning, causing a decrease in the theoretical flow by 82 % in stems and 27 % in petioles. Stem xylem blockage was apparently due to tyloses growing into conduits. We advance the hypothesis that the entire process of leaf shedding and winter rest may be initiated by extensive stem embolism occurring during the summer.     

The prediction of leaf canopy expansion in the leek from a simple model dependent on primordial development

In an analysis of leaf development of leek plants grown in the field in 1988, successive leaves initiated, appeared (tip and ligule) and senesced at equal intervals of accumulated temperature/thermal time. These intervals corresponded to a plastochron of 92°C days and phyllochrons of 135 (tip) and 233 (ligule) °C days. The rate of appearance of ligules was exactly equal to the rate of leaf senescence, with the result that the number of fully-expanded leaves per plant remained constant at 1.4. These data, which were compatible with results from previous seasons, were used to develop a model of the interrelationships between primordium initiation at the shoot apex and subsequent events in the development of individual leaves. Primordium initiation is considered to be the primary controlling event in the life of a leaf, and the processes of tip appearance, ligule appearance and death can be predicted from knowledge of the number of primordia which have been initiated, without reference to the environment. A model of canopy expansion, based on the central role of the shoot apex, was developed using the temperature relations of primordium initiation and additional data on leaf expansion and leaf dimensions. Leaf area indices computed in this way provided a satisfactory simulation of the thermal-time course of leaf area index observed in a previous season, 1985.     

十种园林植物叶片导热系数及对流换热系数测定

为研究植物层传热特性,选取校园内十种常见园林植物测定其叶片导热系数、叶片与周边空气对流换热系数,拟合导热系数与叶片含水量的近似关系式,对比实验测定对流换热系数与通过经验公式理论计算所得对流换热系数,比较与叶片接触前后空气的相对湿度。结果表明,叶片存在降温增湿作用,在5～25 ℃下叶片导热系数随温度变化较小;叶温20 ℃时,叶片导热系数随叶片含水量降低而减小;实验测试对流换热系数与理论计算结果吻合度较高。     

Leaf hydraulics and drought stress: response, recovery and survivorship in four woody temperate plant species

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential ( Ψ _leaf), leaf hydraulic conductance ( K _leaf), and midday transpiration ( E ) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of K _leaf to declining Ψ _leaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and K _leaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of K _leaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to Ψ _leaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery.     

Leaf primordia initiation,leaf emergence and tillering in wheat (Triticum aestivum L.) grown under low-phosphorus conditions

In two simultaneous experiments we examined the effects of phosphorus (P) supply on leaf area development in wheat (Triticum aestivum L.) grown in sand with nutrient solutions. In Experiment 1 we studied leaf emergence, leaf elongation, tiller emergence, shoot growth, and P uptake under four levels of P supply (mM) 0.025 (P1), 0.05 (P2), 0.1 (P3), and 0.5 (P4), and. In Experiment 2 there were two levels of P supply, P1 and P4, and we examined the effects of P on leaf primordia differentiation and leaf emergence. The phyllochron was calculated as the inverse of the rate of leaf emergence calculated from the regression of number of leaf tips (PHY-Ltip), Haun index (PHY-Haun), and as the cumulated thermal time between the emergence of two consecutive leaves (PHYtt). The plastochron was calculated from the inverse of the rate of leaf primordia initiation in the apex. P deficiency delayed the emergence of leaves on the main stem and on the tiller 1. Phosphorus deficiency increased the time from emergence to double ridge and anthesis. The final number of leaves was not affected by P. The effects of P on the value of the phyllochron were attributed to both a reduced rate of leaf primordia initiation, and to a reduced leaf elongation rate. P deficiency delayed or even suppressed the emergence of certain tillers. In this work a phosphorus deficiency that reduced shoot growth by 25% at 44 days after emergence significantly modified the structure of the plants by increasing the value of the phyllochron and delaying tillering. These results suggest that any attempt to simulate leaf area development and growth of wheat plants for P-limited conditions should include the effects of the deficiency on leaf emergence.     

Leaf discs floated on water are different from intact leaves in photosynthesis and photoinhibition

Photoinhibition has been often evaluated with leaf discs floated on water or placed on wet papers to prevent desiccation. Under these conditions, there is a possibility that CO₂ diffusion is blocked by water, which may lead to reduction in photosynthetic CO₂ assimilation. Using Chenopodium album L. grown at two irradiances, photosynthesis, quantum yield of Photosystem II (ΔF/F _m′), non-photochemical quenching (qN), and photoinhibition were compared between detached leaves and leaf discs. In low-light-grown plants, photoinhibition was greater in leaf discs than in detached leaves, while in high-light-grown plants, there was little difference. Leaf discs showed lower rates of photosynthesis and ΔF/F _m′, and higher qN. The ΔF/F _m′ in leaf discs increased when leaf discs were exposed to high concentration of CO₂, suggesting that CO₂ diffusion to chloroplasts was limited in leaf discs floated on water. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthesis and water-relation traits of the summer annual C4 grasses, Eleusine indica and Digitaria adscendens, with contrasting trampling tolerance

Two summer annual C₄ grasses with different trampling susceptibilities were grown as potted plants, and diurnal leaf gas exchange and leaf water potential in each grass were compared. The maximum net photosynthetic rate, leaf conductance and transpiration rate were higher in the trampling-tolerant Eleusine indica (L.) Gaertn. than in trampling sensitive Digitaria adscendens (H. B. K.) Henr. Leaf water potential was much lower in E. indica than in D. adscendens. There were no differences in soil-to-leaf hydraulic conductance and leaf osmotic potential at full turgor as obtained by pressure–volume analysis. However, the bulk modulus of elasticity in cell walls was higher in E. indica leaves than in D. adscendens leaves. This shows that the leaves of E. indica are less elastic. Therefore, the rigid cell walls of E. indica leaves reduced leaf water potential rapidly by decreasing the leaf water content, supporting a high transpiration rate with high leaf conductance. In trampled habitats, such lowering of leaf water potential in E. indica might play a role in water absorption from the compacted soil. In contrast, the ability of D. adscendens to colonize dry habitats such as coastal sand dunes appears to be due to its lower transpiration rate and its higher leaf water potential which is not strongly affected by decreasing leaf water content.     

Acclimation of leaf growth to low water potentials in sunflower

Abstract Leaf growth is one of the most sensitive of plant processes to water deficits and is frequently inhibited in field crops. Plants were acclimated for 2 weeks under a moderate soil water deficit to determine whether the sensitivity of leaf growth could be altered by sustained exposure to low water potentials. Leaf growth under these conditions was less than in the controls because expansion occurred more slowly and for less of the day than in control leaves. However, acclimated leaves were able to grow at leaf water potentials (Ψ₁) low enough to inhibit growth completely in control plants. This ability was associated with osmotic adjustment and maintenance of turgor in the acclimated leaves. Upon rewatering, the growth of acclimated leaves increased but was less than the growth of controls, despite higher concentrations of cell solute and greater turgor in the acclimated leaves than in controls. Therefore, factors other than turgor and osmotic adjustment limited the growth of acclimated leaves at high ψ₁ Four potentially controlling factors were investigated and the results showed that acclimated leaves were less extensible and required more turgor to initiate growth than control leaves. The slow growth of acclimated leaves was not due to a decrease in the water potential gradient for water uptake, although changes in the apparent hydraulic conductivity for water transport could have occurred. It was concluded that leaf growth acclimated to low ψ₁, by adjusting osmotically, and the concomitant maintenance of turgor permitted growth where none otherwise would occur. However, changes in the extensibility of the tissue and the turgor necessary to initiate growth caused generally slow growth in the acclimated leaves.     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.     

Leaf Dynamics of Panicum maximum under Future Climatic Changes

Panicum maximum Jacq. ‘Mombaça’ (C4) was grown in field conditions with sufficient water and nutrients to examine the effects of warming and elevated CO₂ concentrations during the winter. Plants were exposed to either the ambient temperature and regular atmospheric CO₂ (Control); elevated CO₂ (600 ppm, eC); canopy warming (+2°C above regular canopy temperature, eT); or elevated CO₂ and canopy warming (eC+eT). The temperatures and CO₂ in the field were controlled by temperature free-air controlled enhancement (T-FACE) and mini free-air CO₂ enrichment (miniFACE) facilities. The most green, expanding, and expanded leaves and the highest leaf appearance rate (LAR, leaves day^-1) and leaf elongation rate (LER, cm day^-1) were observed under eT. Leaf area and leaf biomass were higher in the eT and eC+eT treatments. The higher LER and LAR without significant differences in the number of senescent leaves could explain why tillers had higher foliage area and leaf biomass in the eT treatment. The eC treatment had the lowest LER and the fewest expanded and green leaves, similar to Control. The inhibitory effect of eC on foliage development in winter was indicated by the fewer green, expanded, and expanding leaves under eC+eT than eT. The stimulatory and inhibitory effects of the eT and eC treatments, respectively, on foliage raised and lowered, respectively, the foliar nitrogen concentration. The inhibition of foliage by eC was confirmed by the eC treatment having the lowest leaf/stem biomass ratio and by the change in leaf biomass-area relationships from linear or exponential growth to rectangular hyperbolic growth under eC. Besides, eC+eT had a synergist effect, speeding up leaf maturation. Therefore, with sufficient water and nutrients in winter, the inhibitory effect of elevated CO₂ on foliage could be partially offset by elevated temperatures and relatively high P. maximum foliage production could be achieved under future climatic change.     

Plant leaf area expansion and assimilate production in wheat (Triticum aestivum L.) growing under low phosphorus conditions

Under phosphorus deficiency reductions in plant leaf area have been attributed to both direct effects of P on the individual leaf expansion rate and to a reduced availability of assimilates for leaf growth. In this work we use experimental and simulation techniques to identify and quantify these processes in wheat plants growing under P-deficient conditions. In a glasshouse experiment we studied the effects of soil P addition (0–138 kg P₂O₅ ha^-1) on tillering, leaf emergence, leaf expansion, plant growth, and leaf photosynthesis of wheat plants (cv. INTA Oasis) that were not water stressed. Plants were grown in pots containing a P-deficient (3 mg P g^-1 soil) sandy soil. Sowing and pots were arranged to simulate a crop stand of 173 plants m^-2. Experimental results were integrated in a simulation model to study the relative importance of each process in determining the plant leaf area during vegetative stages of wheat. Phosphorus deficiency significantly reduced plant leaf area and dry weight production. Under P-deficient conditions the phyllochron (PHY) was increased up to a 32%, compared to that of high-P plants. In low-P plants the rate of individual leaf area expansion during the quasi-linear phase of leaf expansion (LER) was significantly reduced. The effect of P deficiency on LER was the main determinant of the final size of the individual leaves. In recently expanded leaves phosphorus deficiency reduced the photosynthesis rate per unit leaf area at high radiation (AMAX), up to 57%. Relative values of AMAX showed an hyperbolic relationship with leaf P% saturating at 0.27%. Relative values of the tillering rate showed an hyperbolic relationship with the shoot P% saturating at values above 0.38%. The value of LER was not related to the concentration of P in leaves or shoots. A morphogenetic model of leaf area development and growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and to study the sensitivity of different model variables to changes in model parameters. Simulation results indicated that under mild P stress conditions up to 80% of the observed reduction in plant leaf area was due to the effects of P deficiency on leaf emergence and tillering. Under extreme P-deficient conditions the simulation model failed to explain the experimental results indicating that other factors not taken into account by the model, i.e. direct effects of P on leaf expansion, must have been active. Possible mechanisms of action of the direct effects of P on individual leaf expansion are discussed in this work.