Canopy structure, light microclimate and leaf gas exchange of Quercus coccifera L. in a Portuguese macchia: measurements in different canopy layers and simulations with a canopy model

SummaryThe structural characteristics of a diverse array of Quercus coccifera canopies were assessed and related to measured and computed light attenuation, proportion of sunlit foliage, foliage temperatures, and photosynthesis and diffusive conductance behavior in different canopy layers. A canopy model incorporating all components of shortwave and longwave radiation, and the energy balance, conductance, and CO2 and H2O exchanges of all leaf layers was developed and compared with measurements of microclimate and gas exchange in canopies in four seasons of the year. In the denser canopies with a leaf area index (LAI) greater than 5, there is little sunlit foliage and the diffuse radiation (400–700 nm) is attenuated to 5% or less of the global radiation (400–700 nm) incident on the top of the canopy. Foliage of this species is nonrandomly distributed with respect to azimuth angle, and within each canopy layer, foliage azimuth and inclination angles are correlated. A detailed version of the model which computed radiation interception and photosynthetic light harvesting according to these nonrandom distributions indicated little difference in whole-canopy gas exchange from calculations of the normal model, which assumes random azimuth orientation. The contributions of different leaf layers to canopy gas exchange are not only a function of the canopy microclimate, but also the degree to which leaves in the lower layers of the canopy exhibit more shade-leaf characteristics, such as low photosynthetic and respiratory capacity and maximal conductance. On cloudless days, the majority of the foliage in a canopy of 5.4 LAI is shaded —70%–90% depending on the time of year. Yet, the shaded foliage under these conditions is calculated to contribute only about one-third of the canopy carbon gain. This contribution is about the same as that of the upper 13% of the canopy foliage. Computed annual whole-canopy carbon gain and water use are, respectively, 60% and 100% greater for a canopy of 5 LAI than for one of 2 LAI. Canopy water-use efficiency is correspondingly less for the canopy of 5 LAI than for that of 2 LAI, but most of this difference is apparent during the cool months of the year, when moisture is more abundant.

[1]  J. Tenhunen,et al.  Limitations due to water stress on leaf net photosynthesis of Quercus coccifera in the Portuguese evergreen scrub , 1985, Oecologia.

[2]  J. Tenhunen,et al.  Development of a photosynthesis model with an emphasis on ecological applications , 1979, Oecologia.

[3]  Margaret C. Anderson Stand Structure and Light Penetration. II. A Theoretical Analysis , 1966 .

[4]  O. Correia,et al.  Structure and dynamics of Serra da Arrábida mediterranean vegetation , 1982 .

[5]  J. Tenhunen Plant response to stress : functional analysis in Mediterranean ecosystems , 1987 .

[6]  H. Mooney,et al.  The energy balance of leaves of the evergreen desert shrub Atriplex hymenelytra , 1977, Oecologia.

[7]  J. Wilson,et al.  INCLINED POINT QUADRATS , 1960 .

[8]  J. Burt,et al.  EFFECT OF RECEIVER ORIENTATION ON ERYTHEMA DOSE1,2 , 1979 .

[9]  J. Tenhunen,et al.  Method for Field Measurements of CO2-Exchange. The Diurnal Changes in Net Photosynthesis and Photosynthetic Capacity of Lichens under Mediterranean Climatic Conditions , 1985 .

[10]  J. Tenhunen,et al.  Effects of temperature at constant air dew point on leaf carboxylation efficiency and CO2 compensation point of different leaf types , 1985, Planta.

[11]  B. E. Mahall,et al.  Drought and changes in leaf orientation for two California chaparral shrubs: Ceanothus megacarpus and Ceanothus crassifolius , 1985, Oecologia.

[12]  J. Tenhunen,et al.  Midday Stomatal Closure in Mediterranean Type Sclerophylls under Simulated Habitat Conditions in an Environmental Chamber , 1982 .

[13]  R. J. List Smithsonian Meteorological Tables , 2018, Nature.

[14]  C. T. Wit Photosynthesis of leaf canopies , 1965 .

[15]  J. Ross The radiation regime and architecture of plant stands , 1981, Tasks for vegetation sciences 3.

[16]  W. Brutsaert On a derivable formula for long-wave radiation from clear skies , 1975 .

[17]  P. G. Jarvis,et al.  Productivity of temperate de-ciduous and evergreen forests , 1983 .

[18]  S. Rambal,et al.  Changes in aboveground structure and resistances to water uptake in Quercus coccifera along a rainfall gradient , 1987 .

[19]  J. Ehleringer,et al.  Leaf absorptance and leaf angle: mechanisms for stress avoidance , 1987 .

[20]  W. Oechel,et al.  Energy utilization and carbon metabolism in mediterranean scrub vegetation of Chile and California , 1979, Oecologia.

[21]  J. Norman,et al.  Photosynthesis in Sitka spruce (Picea sitchensis (Bong.) Carr.). V. Radiation penetration theory and a test case , 1975 .

[22]  A. E. Hall,et al.  Stomatal Responses, Water Loss and CO2 Assimilation Rates of Plants in Contrasting Environments , 1982 .

[23]  R. W. Pearcy,et al.  Photosynthetic Responses to Dynamic Light Environments by Hawaiian Trees : Time Course of CO(2) Uptake and Carbon Gain during Sunflecks. , 1985, Plant physiology.

[24]  P. Miller,et al.  BIOCLIMATE, LEAF TEMPERATURE, AND PRIMARY PRODUCTION IN RED MANGROVE CANOPIES IN SOUTH FLORIDA' , 1972 .

[25]  P. Jarvis,et al.  Modelling Canopy Exchanges of Water Vapor and Carbon Dioxide in Coniferous Forest Plantations , 1985 .

[26]  James R. Ehleringer,et al.  Non‐random leaf orientation in Lactuca serriola L. , 1984 .

[27]  H. Mooney,et al.  Leaf age and seasonal effects on light, water, and nitrogen use efficiency in a California shrub , 1983, Oecologia.

[28]  William A. Williams,et al.  A model for simulating photosynthesis in plant communities , 1967 .

[29]  J. Tenhunen,et al.  Midday stomatal closure in Mediterranean type sclerophylls under simulated habitat conditions in an environmental chamber , 1981, Oecologia.

[30]  D. N. Baker,et al.  Simulation of Growth and Yield in Cotton: I. Gross Photosynthesis, Respiration, and Growth 1 , 1972 .

[31]  J. Ehleringer,et al.  Leaf absorptances of Mohave and Sonoran desert plants , 1981, Oecologia.

[32]  I. R. Cowan The Interception and Absorption of Radiation in Plant Stands , 1968 .

[33]  C. Field Leaf Age Effects on the Carbon Gain of Individual Leaves in Relation to Microsite , 1981 .

[34]  W. E. Williams Optimal water‐use efficiency in a California shrub , 1983 .

[35]  J. Tenhunen,et al.  The control by atmospheric factors and water stress of midday stomatal closure in Arbutus unedo growing in a natural macchia , 1982, Oecologia.

[36]  P. Miller,et al.  Canopy Structure and Environmental Interactions , 1979 .

[37]  S. Rambal,et al.  Water balance and pattern of root water uptake by a Quercus coccifera L. evergreen srub , 1984, Oecologia.

[38]  I. R. Cowan,et al.  Stomatal function in relation to leaf metabolism and environment. , 1977, Symposia of the Society for Experimental Biology.

[39]  John M. Norman,et al.  Light Intensity and Sunfleck‐size Distributions in Plant Canopies1 , 1971 .

[40]  J. Tenhunen,et al.  Diurnal Courses of Stomatal Resistance and Transpiration of Wild and Cultivated Mediterranean Perennials at the End of the Summer Dry Season in Portugal , 1982 .

[41]  S. Rambal Water balance and pattern of root water uptake by a Quercus coccifera L . evergreen scrub , 2004 .

[42]  W. A. Stoner,et al.  Models of Plant and Soil Processes , 1981 .

[43]  J. Monteith Light Distribution and Photosynthesis in Field Crops , 1965 .

[44]  E. Schulze,et al.  A portable steady-state porometer for measuring the carbon dioxide and water vapour exchanges of leaves under natural conditions , 2004, Oecologia.

[45]  D. Baldocchi,et al.  Seasonal variation in the statistics of photosynthetically active radiation penetration in an oak-hickory forest , 1986 .

[46]  B. Juniper THE EFFECT OF PRE‐EMERGENT TREATMENT OF PEAS WITH TRICHLORACETIC ACID ON THE SUB‐MICROSCOPIC STRUCTURE OF THE LEAF SURFACE , 1959 .

[47]  O. Lange,et al.  Photosynthese und Wasserhaushalt der immergrünen mediterranen Hartlaubpflanze Arbutus unedo L. im Jahreslauf am Freilandstandort in Portugal: I. Tagesläufe von CO2-Gaswechsel und Transpiration unter natärlichen Bedingungen , 1986 .

[48]  M. Monsi Uber den Lichtfaktor in den Pflanzengesellschaften und seine Bedeutung fur die Stoffproduktion , 1953 .