TBl..AU) TO rIOm:L GBOWTR OJ' BUCX\il1EA!r PLAB'T CABOPI FROn DINAlUC
POIlfT OF VIEW Xaj!el-Bogataj and Andrej Univerza Edvard.a Kardelja T' Ljubljani, lHotehniska !altulteta, VTOZD za agronomijo, Ljubljana, JugoslaVijal) A dynamic multi la;yered plant canopy model llUCKPRO based on lea! temperature and irradiance is presented. It simulates the day to day assimilation of photosynthetic material. Results are presented and compared with observed productivity. IBTBODUCTIOH Growth and development of planta depend on their environment. Their interrelations are studied with tva different groups of methods, viz. empirical-statistical ones which originate from local conditions and usually can not be generalised and dynamic ones which have this important property. Dynamic methods usually named modele simulate day to d&7 assimilation on the basis of energy and mass '._-------- l)University Edvard Kardelj of Ljubljana, Department of Agronomy Ljubljana, Yugoslavia - ,0 - exchange among the plants and their environment and among various processes taking place in the plants themselves. Both groups ot, methods deliver useful results but the last one should be pretered being based on physical, chemical and biological principles. In our york ve develop a dynamic model BUCKPBO as a computer program. It simulates buckwheat canopy netphotosynthesis on the basis ot input data, viz. meteorological data, ,buckwheat pllQlt " stand architecture and physiological characteristios. Serious problem was the lack of adequate data for the buckwhea:t plant canopy. It was solved partly with our own measurements a j f e ~ Bogataj, Knave 1985 and KajfeZ:.-Bogataj et a1. 1986) and com- pleted from the literature (Evans 1978, Horie 1980, Hesketh 1980, r.;onteith 1975, Akberdina 1970). THE BUCKPBO MODEL BUCKPRO is a multi-layered buckwheat canopy model derived trom CALPBOD model (Terjung 1981, Band et ale 1981, Kajfez- Bogataj and Robie 1985). It ca.lculates primary net assimilation (P) on the basis of leaf temperature (TL) and its irradiance (SGR) using appropriate plant photosynthesis response curves: LE L - )1 - The basic equation for the energy budget ot a leaf from which lea! temperature is calculated can be written algebraically: where as - shortwave lea! absorptivity, aL - longvave leal absorptivity, r - groulXl albedo (F lot- - incident longvave radiation aoove lea! (w/m 2 ), It - incident longvave radiation below lea! (lJ/ m 2 ), B GR global radiation (V/m 2 ) t L - lea! emissivity, G - Btetan-Boltman constant (lJ/m 2 r;!1-), TL -lea! temperature LE - latent heat flux density (w/m 2 ), L - latent heat of vaporization (J/kg), R - sensible heat flux density (W/m 2 ). The shortwave components of Eq. (2) can be derived by a ot methods (Hocevar et ale 1982) tor unobstructed, horizontal areas and several methods exist tor the simulation of longwave radiation, as well (Bruntsaert, 1975). The equivalent ot lea! transpiration is given as and Papian, 1971): Af , - 32 - where: A f - ti!!erence between saturation vapor _onsities at lea! and air tempera'ture, r a - boundary layer resistance to vater dit'!usion (s!cm) , r s - stomatal resistance to water vapor cliffusion (s!cm). Boundary layer resista.ucei.s calculated .trom lea! dimension, lea.!' wiclth and wind velocity. Stoinatal resistance vas aBSUlIled as a function only of lea! temperature ranging .trom a maximum at oOe to a minimUm at opti.mum temperature (TOPT). The last term in Eq. (2) calculates tho sensible heat flux of the leaf: H. h bT c where h c - convection coer.ticient, A T - difference between lea! and. air Convection coefficient (he) also depends on leaf dimensions and rind. ve1oeity Shortwave radiation arriving at a canopy level z on a horizontal plane SGH(z) dimishes exponentially with increasing path: - )) - where EGa (top) global radiation at the top of the canopy, x - radiation extinction coefficient, LAI(Z) - leaf index Values of EGR(Z) are then used for calculation of TL (eq.(2 at different heights ana for calculation of netphotosyntheeis via plant response curves. The model assumes optimal vater supplies, fertilizer applications and disease control (Pig. 1) but can be addapted to high evaporative demand.s by changing input parameters - for example stomatal resistance. Photosynthesis was then modeled 8S a reaction to different lea! temperatures via a sistem of photosyntetic response curves, (Fig. 2). - 34 - INPUT ,llTREME DULY ;Air tecperature I jir'i.nd velocity ~ l humidity ! , I ! i ! I i liEAH DAILY, OTHER Declination Rad.ius vector Latitude Solar constant Elevation Daily solar radiation Cloudiness DABOPI-LEAF VARIABlES FOR BUCK'JHEA.T Leaf d.imensions Upper threshold temperature Optimum lea! temperature Minimum stomatal resistance Poliage albedo Foliage emissivity Leaf transmisivity Soil albedo S o ~ l conductivity Radiation extinction coefficient Lea! area index Stomatal resistances Bound.ary layer resistance Convection coefficient for all layers Attenuation and exchanges of shortwave radiation Attenuation and. exchanges of longwave radiation IEnergy budget I of leaf layers Iiterationsl Leaf temperatures for all layers OUTPUT ' PHOTOSYNTHESIS Net photosynthesis, Sum of stand Cumulative sum of stand netphotosynthesis I from response curvesl rig. 1: Generalized BUCKPRO flow Chart, emphasiZing input and output. - :55 - 8 - :.c '1
'1J ,.., , 6 0 5<3R '" VSW/rfi u 01 5 oS tf) m 0 /. w :J: 0 t- 3 z lj) 0 2 t- o 0 it 0 LEAF TEMPERAlURE ("C1 0 1) '6 20 25 30 40 Fig. 2: Comparison of observed data and approximative net photosynthesis response curve for Fagopyrum esculentam Moench (PAR = 135 Wm- 2 , SGR = 275 Wm- 2 ) We estimated net photosynthesis response curves [0= buckwheat (Fagopyrum esculentum Moench) from ty Infra red gas analyser (IGRA, The Analytical Co. Ltd., Type 225/2, Hoddeson, England) in climate chamber (Gorenje, TGO, Titovo Velenje, Yugoslavia) as a function leaf temperatures, measured by tele-termometers (YSI, Model 44 TZ, Yellow Ohio, USA) and artificial PAR radiation measured by (LI-COR, LI 1985). Frou. cata - 36 - we derived a family of approximative anallytical curves for various values of SGR which were used in the model. INPtJT CONSIDERA!rIOliS The model uses three groups of input data astronomical (solar radius vector, solar declination, latitude, elevation, solar constant), meteorological (minimum and maximum daily air temperature, relative humidity, wind speed, solar radiation and cloud cover)and biological ones. Function generators were used to interpolate hourly data from minimum and maximum values. curves are determined for air temperature, wind speed, relative humidity and the positive half of a sine curve for hourly values of shortwave radiation. In our case we used decade daily data from August 1at 1984 till October loth for Ljubljana. meteorological data the air temperature and the solar radiation have the influence on the results of the model. More troubles caused third group of input data determinning bu:rwheat stand and lea! variables together with soil characte- ristics (Pig. 1). The typical lea! dimension for buckwheat is 5 cm, the upper threshold temperature is 55C, optimal tempera- ture accordins to our measurments (Fig. 2) was 20C. According to (Larcher 1980, Ross 1981, Sivakumar 1984, Terjung and 1950) the foliage albedo used was 0.1, foliage - J7 - emissivity 0.96, the radiation extinction coefficient 1, soil albedo 0.15, soil conductivity 0.7 transmissivity 0.2 and minimal stomatal .resistance ; cms- l
The most important parameters in this group are the leaf
area index and minimal stomatal resistance. Especially lea! area index changes during the vegetation period. We studied this parameter during buckwheat development in 1984 (Kajfez-Bogataj, lCnavs 1985). Lear areas vere measured by automatic planimeter type LI-COR (LI ;050 A) for each ot 5 stages ot stand development (Table 1) and the results vere used as input to the model. Table 1: Top and bottom lqer contributions to cumulative -LA! during the growth period ot buckwheat, sown on AUgust 1at, 1984 in LjUbljana dqe .'iltter sowing 18 ;2 46 60 74 L A I top half 2.8 4 2.6 4.5 bottom half 1.8 3.6 0.9 0.8 CUMULATIVE LA! 4.5 4.6 7.6 ;.5 2.1 - J8 - RESULTS AND DISCUSSIONS By means of the described model net vas sLmulated using mean decade daily data for the vegetation period August 1st - October loth, 1984 tor Ljubljana (300 m.L\.reather condi tiona during that period vere not optimal, lov temperatures and high precipitations vere characteristic tor the tirst halt of .August and for the second halt ot: September. Dry production vas therefore belov expectation. The calculated values vere compared with observed data vhich vere obtained on the basis of the regular sampling ot buckwheat dry matter production between tvo consecutive dates. F1gure 3 shows a comparison between calculated and measured values of dry matter production. Both values correspond quite vell though calculated values are higher than the observed ones. Calculated total dry matter production is about 12 %higer than the observed one. - 39 - LJUBLJANA - 1984 0 ,/ .s::. / -. / ,/
/ W f l- I- I eo <l: I > / cc I 0 / 60 ..J I <f I 0 I I- / ---- CALCUlATED 4 / / / OBSERVED / 20 I / / / I nAYS AFTER SOWING 10 20 30 JIJ 50 60 70 160 Fig. 3: Comparison of observed and simulated buckwheat total dry matter production for.Ljubljana in 1984 - 40 - The reasons vhy the calculated and observed data do not mach can be traced in the folloving: - Some important informations about experiJnental data are incomplete(optical properties of stand, stomatal resistances etc) - The present knowledge of buckwheat reactions to the environment is rather ucleared especially regarding storage and redistribution of the produced assiJnilates. - Experimental data obtained in the climatic chamber were transfe- red to the ambient condi tiona. This assumption is uncertain and needs testing. - Rough observations of biomas production {the ones used did not include root mass and senesence) and ~ enough good estimatee of meteorological elements at the field site studied (now data are from metecrological station about 10 km away) contributed to discrepancy. To obtain a better agreement between the results of the model and observed productivity of buckwheat stand the further research should be concentrated on the mentioned problems. - 41 - Beside presented results the model has numerous options regarding experimental sites, atmospheric input and biological characteristics. Weather data can be for specific years, for long term climatic averages or tor prescribed (for instance extreme) data, depending on user s objectives. nodel can be applied also to various genotypes and buckwheat species by using their physiological and plant architecture characteristics. CONCLUSIONS The netphotosynthesis model BUCKPRO of buckwheat stand productivity as a function of environment is presented. Essential input data concerrJUlg buckwheat physiological and stand characteristies were measured and disseused. A fairly good agreement between the simulated and observed values of netphotosynthesis was obtained. The possible couses of disagreement are disscused together with the options of the model. be made to cbtain more acurate data on stand parameters (stomatal resistance, optical properties, tAr) during the growing period. Such data essential to any dynamic modelling of plant growth and productivity. - 42- .A.CDO'WLEDGEME!lT We are 'gratetul to DJ: ... J.. 1'1arlincic and !'lag. A. Gabricek' tor their assistance. REP'ElmJCES 1 .1kberdina R.H. (1970). Vlijanie temperatun vozduha na dnevnoi hod intenzivnosti !otosinteza i intensivnosti ekzosJIlOsa vodi iz listev grecihi. Selekcija i agrotehnika greHhi, VllIIZX, Orel, 65-72. 2. Band et &1. (1981). Application ot a photos;ynthetic&1 model to an agricultural region ot varied climate$: California. Agric. Metoorol. 24, 3, 201-218. 3. Evans, L.T. (editor) (1978). Crop physiology, Cambridge University Press, Cambridge) 374 pp. 4. Hocevar .l. et all. (1982). Boneno obsevanjo v Sloveniji, VDO BP', VTOZD za agronomijo, Zbornik BP', supl. 6, 97 pp. 5. Hocevar, .l., Xajtez-Bogataj, L. (1984) .lplikacija modela neto!otosintezo na klimatsko razlicnih obmocjih Slovenije, Zbornik BP' Univerze E.K. v Ljubljani"43, 9 3 ~ - 4) - 6. Horie, T. (1980). Photos;ynthesia and Primary Production Rice Plants, Journal of .A.gricultural Meteorology, The Societ)" of Agricultural Meteorology of Japan, Vol. 35, Ho. 4, 201-215. 7. J.D., (1980). Predicting photos;ynthesis for ecosystem models, Vol. II., CBS Press, Boca Raton, Plorida, 279 pp. 8. L. and Knavs 11. (1985). Studies on the production of dr7 matter in the community of buckwheat with particular reference to lea! area. Pagopyrum (Ljubljana), Vol. 5, 7-13. 9. Xajfel-Bogataj L. and D. Comparison of observed and predicted forest productivity in various climatic conditions in Slovenia, .Seminar on biometerological methods, proceedings, Zbornik Bl,' supplement 10, 57-67. 10. I:.djfez-Bogataj L. et ale (1986). Analysis of different net photos;ynthesis response curves for buckwheat. Pagopyrum, Vol.6 (in 'preparation) 11. Larcher W. (1980). Physiological Plant Ecology, Springer- Verlag,' Hew York, 304 pp. 12. Ross, J., (1981). The radiation regime and architecture of plant at ands, W. Junk Publishers, London 1981, 391 pp. - 44 - 13. J.L., (1975). Vegetation and the Atmosphere, Vol. 1., Principles, Academic Press, London, 2?8 pp. 14. Sivakumar, 1'1., (1984). Crop productivity in rela:tion to interception of photosynthetically active radiation. Forest and agricultural meteorology, Vol. 31, No.2, 131-14;. 15. Terjung, W.li , O'Rourke, (1980). An Economical Model for use in Urban CliJnatology, Int. J. Biometeoro1ogy, Vol. 24., no. 4, 281-291.
Photosynthetic Activity of Intertidal Microphytobenthic Communities During Emersion: in Situ Measurements of Chlorophyll Fluorescence (Pam) and Co Flux (Irga)