Acta cologica, 1992, 13 (5), $35-542 Net photosynthesis-leaf temperature relations in plant species with different height along an altitudinal gradient Rada F. (*), J. Gonzalez (**), A. Azocar (*), B. Bricefio (*) and R. Jaimez (***) (9) Contra de FvestigacomesEclogics de los Andes Tropicals (CIELAT), ‘acted de Cencas, Unersiad dels Andes, Merida, Venue (4) Fundacion Miguel Lil, Tucunan, Argentina, (°°) Ctr de Simulacion y Models (CESIMO), Universidad de ls Andes, Meid, Vena Abstract ‘The relationship between net photosynthesis and leaf temperature was studied along a gradient inthe topical high Andes in order to determine if there were any diferences in CO assimilation parameters due to temperature changes along the Solsit continuum, Two acaulescent rosetes, Calandrinia acaulis and Hypochoers setosus, shrub, Hypericum laricodes, and a gia ovet, Expeleia schuli, were chosen for this study at altudes from 2950 to 4 200m. Optimum lemperature for photosynthesis decreased from 115°C to 7.5°C for E,schultei and from 139°C {© 83°C for H.laicodes at 2.950 and 4 000 m, respectively. H. Seiosus showed an increase in ‘optimum lea temperature for photosynthesis from 3 $500 4200 m (8.8°C and I1.1°C, respectively, and C. acauls showed no diflerences between aitudes (approximately 130°C), Net photosynthetic fates were also higher forthe two ground level plants (65 to 8.0 jamal mr?!) compared to the higher level plans (4.1 t0 4.6 jzmol me"), All species showed postive assimilation rates at Below °C, Optimum temperature for photosyhesis was coupled to ai temperatures forthe high stratum species, therefore decreasing at higher ahitude. The lower stratum species, strongly influenced by soil temperature, showed similar or even an increase in optimam lea temperature for photosynthesis ‘with an inerease in alitade ‘Keywords: Net photosymtbesis, laf temperature, life forms, high tropical mountains, Résumé {Les relations entre la photosynthise mete et la température des feuiles ont été Gudies le long un gradient dans les hautes Andes tropicals ain de détemniner sl y a des differences dan les aramécesd'asimilation du CO; dies des changements de temperature au lng da continuum pol sir. Deux rosettes saulescentes,Calandriniaacaulis et Hypochoers setosus, wn arbuste,Hyperieurt laricoides et une rosete ante, Espeeraschuli ont 6 choisis pour cette ute, a des atts allant de 2.950 m i-4 200 m. La temperature optimim pour la photosynth décroit de 11,5°C 3 15°C pour E.schulal et de 139°C A 8.3°C pour H.larcoides, 42950 m et 4 000m respectivement 2H serous présente une augmentation de la température des files optimim pour la photosynhise, (4 3580 m 24200 m (88°C et 11,7°C respectivement) et C.acauls ne présente pas de diérence entre les altitudes (approximativement 13°C), Les taux nets de photosynthese sn Gealement pls Aca Beologica 1146-609 XP205/S35908S 280/ © Gauthis-Villar536 F. Rada ef al loves pour les deux planes basses (65 88.0 aol ms!) que pour les deux planes plus havtes (4.1846 pmol mrs"). Toutes les espices préSentent des taux assimilation positifsau-dessous de (0°C, La temperature optimum pour la photosynthse est ie ala empératre desir pour ls planes Inautes, elle déeroit dane & des alas pls élevés, Les planes basses, forementineencées par temperature do so, présente une température optimum pour la photosyathese similsire ou méme plus elevee avee une augmentation de altitude INTRODUCTION Plant morphology strongly influences the day-time temperature regime of plants (Noss, 1988). Typical life-forms for tropical alpine environments have been described by many authors: giant rosettes are one of the most important cones (Espeletia in the Andes; Custmecasss, 1979; Swrm, 1981; Dendrosenecio and Lobelia in East Africa; Heoseso, 1964; Cos, 1967). Together with these giant rosettes, vascular plants may be included into four conspicuous growth forms: 1) tussock graminoids, 2) dwarf shrubs, 3) herbaceous, mostly perennial rosette plants and 4) cushion plants (Sam & Youo, 1987; Kowver & Laxcier, 1988). The first two forms are plants that grow well above ground level, while the last two grow at most a couple of centimeters above ground. AA temperature gradient exists along soil-air profiles, day-time air temperature closer to the ground are always significantly higher than those further away from the ground (Movastenio, 1979; Azocar & Moxastenio, 1980) which means that those plants growing closest to the ground will be subjected to greater daily leaf temperatures. Low plant canopies in alpine regions are substantially warmer during the day but cooler during the night with respect to air temperature (Konwen & Laxcuer, 1988). With respect to the colder nights near the ground for the tropical Andes, Azocak et al. (1988) and Soueo ef al. (1991) have found that plants near the ground use mainiy tolerance mechanisms to survive under these more extreme ow temperature conditions, while those plants that grow above ground level use avoidance mechanisms, mainly through supercooling, to resist the less rigorous night-time conditions (Rapa et al., 1985; Gotosren et al, 1985). On the other hand, the higher day-time temperatures must have different effects on the ground vegetation with respect to the taller growing plants Since leaf temperature is one of the main determinants of the photosynthetic capacity of plants we would expect to find differences in certain parameters such as ‘optimum leaf temperature for photosynthesis and temperature compensation points for plants growing in different strata. Therefore, the purpose of this work was to study how this difference in leaf temperature influences the photosynthetic capacity of plants with different height, comparing on one hand two acaulescent rosettes, and on the other a small shrub and a giant rosette MATERIALS AND METHODS Four diferent species from two different aliudes were chosen fortis study: Calandriniaacauls (Pomulacaceae) and Hypochoeris setosus (Asteraceae), two acaulescent rosettes growing at 3 $50 m (Paramo. de Mucubaj) and 4 200 m (Paramo de Piedras Blancas) were selected as the ground level Plants, Both species have thin membranaceous leaves, the forme 3 to 6 em long and Oo 0.7 em wide And the later 3 t 6 em long and I 10 1.5 em wide. Espeletiaschultsii (Asteraceae), a giant eavlescent Acta Gcologice [Net photosgnthess ~ Leaf temperatur rosete, and Hypericum laricoides (Hy and 4 000 m (Pio El Aguila) were the leaves 20 to 40 em long and 3 105 em approximately 0.5 em long. Addition eserbed in Mosasrbuo de RevEs (19 Several plants (n=4-5) ofeach sp soil from the study site, transported t ht (12 h photoperiod, 600 jimmy ight period, 5°C dak period at 2 950 period, 0°C dark period at 4 000 and & fut in the laboratory using a gas exch and a plant chamber (O14 m'). The cun laboratory completing all measurement hours. The chamber was equipped wit 8 chart recorder to aban iran lat also fited into the chamber for light me the chamber. A refrigerated bath ith ai temperatures and consequently leaf measured through CO} depletion from to the chamber environment. These at each chosen temperature (approxima ‘emperatures near the optimum). Plant Photosynthesis (A: mol mS") wak A where: V isthe chamber volume (en), (umol mv) per time (seconds) (Fi3 RESULTS Leaf temperature-net photo: schultz and Hypericum laricoi temperature for photosynthesis 4.000 m, and for Hypericum 83°C at 4.000 m. Maximum p 4.0 and 5.0 mol ms". Figur relations in the lower stratum sp As opposed to the higher strat temperature optimum with incr. and 11.7°C, respectively). In in leaf temperature optimum t [Net photosynthetic rates for the ‘compared to the high stratum s includes mean values and stands each species at each altitude, Th comparing different altitudes for C. acaulis does not show any sig three species show differences ir temperature compensation point Vol 18, n° 5 - 1992of have ‘ant giant 1) slants. first lgrow ature from } that y leaf turing jen & bpical near jreme Huse ‘orous Nother kound ‘thetic ich as points fas to pacity rettes, ‘acaulis 550 m ud level rm wide slescent bologica ‘Nat photosymthests ~ Lea temperature relations 337 roe, and Hypericum laricoides (Hypercaceae), «small shrub; growing at 2.950 m (Las Plants) "Aguil) were the two higher stratum plants. E schulai has very thick pubescent lees 20 1040 om long and 3 t0 5 cm wide; while H.laricoides has membranaceous sale-lke leaves ‘pproximately 0.5 em long. Additional information on site descriptions and climatic conditions are dtsenbed in Moxasteho & REVS (1980), Go.ssTe\ etal. (1985) and Rana e al. (1987), Sever plants (n=4-5) ofeach species at each shtude were excavated with rots and surrounding sol fom the stagy site, ranspored to the Ieboratory and placed in a growih chamber equipped with Jak (12h photoperiod, 600 jamol ms!) and temperature contol simulating Geld conditions (15°C lik period, S®C dark period at 2 950 m; 12°C light period, 3°C dark period ar 3 580 m; 7°C light rod °C dark period at 4 00 and 4 200 mm). Net photosynthesis Taf temperature curves were carried ‘inthe laboratory using a gas exchange system with an infrared gas analyzer (LCA-2, ADC Ltd.) tnd a plant chamber (014 m’). The curves were started immediatly aftr the plans were brought the laboratory completing all measurements fora speccs ta given altitude ina period no longer than 30, ‘nurs. The chamber was equipped with three 36-gauge copper-constntan thermocouples connected ‘char recorder to obtain ar and lea temperatures. A Photosyatheially Active Radiation sensor was ‘to fted ito the chamber for ight measurements. A 1 000 mol ms" light source was placed above the chamber, A refrigerated bath with hose connection to radiator was used to increase or decrease si temperatures and consequently leaf temperatures within the chamber. CO2 assimilation rates were ‘mesured through CO? depletion from the chamber making sure only photosynthetic tissue was exposed tothe chamber environment. These measurements were dane at I minute intervals for five minutes seach chosen temperature (approximately 3°C intervals at high and low extremes and 0.5 to 1°C at Temperatures near the optimum). Plants were left fom 10 to 15 minutes at each set temperature. Net photosynthesis (A: mol ms") was calculated through: A=V+ACOo/L» Atime bere: V isthe chamber volume (mn), 1s he leaf area (mn), NCO; are changes in COs concentrations (amot mr’) per time (seconds) (FID e a, 1989). RESULTS Leaf temperature-net photosynthesis relations for the two taller plants, Espeleria schulizi and Hypericum laricoides, ae shown in figure 1. For E. schultzii optimum temperature for photosynthesis decreased from 11.5°C at 2 950 m to 7.9°C at 4000 m, and for Hypericum laricoides it decreased from 13.9°C at 2 950 m to 83°C at 4 000 m, Maximum photosynthetic rates for both species were between 40 and 5.0 mol ms". Figure 2 shows the leaf temperature-net photosynthesis lations in the lower stratum species, Calandrinia acaulis and Hypochoeris setosus As opposed to the higher stratum species, H. setosus showed an increase in leaf temperature optimum with increase in altitude from 3 550 m to 4 200 m (8.8°C and 11.7°C, respectively). In the case of C. acaulis, there were no differences, in leaf temperature optimum between the two altitudes (approximately 13°C Net photosynthetic rates for these lower stratum species were significantly higher compared to the high stratum species (between 6 and 8.5 jumol ms"), Table I includes mean values and standard errors for the different measured parameters for each species at each altitude. There are no differences in term of net photosynthesis comparing different altitudes for each species. With respect to the other parameters, C.acaulis does not show any significant differences in any of them, while the other tire species show differences in optimum temperature for photosynthesis and high femperature compensation point. In relation (0 the low temperature compensation Wa 13, n° 5 = 192538, F, Rada et al scanty anions | | Fo. 1. — Net photsyntesisleaf temperature cures forthe, two high stratum plants at two dierent ‘ude. Espeleia sch: 4) 2 980 m, y = -O012+0.257%+2878, 7-89; and b) 4 O00 m, Y= -O012x!40.1804+2.950, = 91, Hypericum laricoides: ¢) 2 980 m, y= -O.COIx+0.243x62.60, FB; and d) 4 000 m, y 0.024840 397%+2.60, P=78, i We ALAS a | : i | 2 BS a IGE peek Fao, 2. ~ Net photosynthesis leaf temperature curves forthe 1wo high stratum plans at wo lilt aittudes. Calandrinia acai: a) 3 580 m, y = ~D.021W*40.590xt4.10, 2.78; and b) 4 200 m, y= 00173%s0.465x08.49, abl. Hypochoerisseosus: €) 3 $50 m, y = -O032x%40.580%+283, Pat; and d) 4 200 m, 9 = -0.025x°+0.6894215, 75:82, point only H. laricoides shows significant differences. Two important results stand Dut: in the first place, optimum temperature for photosynthesis are much lower in the higher stratum plants as compared to the ground level ones. And secondly, net photosynthetic rates calculated on a leaf area basis are much higher in the ground level plants Acta eoogica [Net photosynthesis ~ Leaf temperat ‘Tsou 1. Low temperature compensa ‘optimum temperature or photos fiw studied species at diferent Species Alte E.schull 2950 m +4000 m H.larcoides 2950 m +4000 3550 m 4200 m 1H seasus 3380 m +200 m {OPT FOR PHOTOSYNTHESIS ws FR. 3. Optimum temperature for Espeletia schulei (@), Hypericum lori ‘The high temperature con 4.000 m, this temperature. fo and 25°C for E. schultzit. Wh ‘compensation points at much hi . acaulis, respectively (figure compensation points there are low temperature. compensation while the other three species h points with approximately ~3.0 ‘The differences in optimum: chosen species are therefore w (Gg. 3). The two higher strata sp optimum for photosynthesis wi the acaulescent species, 7. ser acaulis maintains similar Value DISCUSSION Many authors have describ high temperatures compared to leaf temperatures of 8 to 10°C the Rocky mountains. Sauisavky Val 18, nt 5 = 1992Net photosynthesis ~ Leaf temperature relations 539 Taste L— Low temperature compensation point (LTCP), high temperature compensation point (HTCP), ‘optima temperature for photosynthesis (P (op) and maximum ne phoosyuhess (Ps (mas) forthe four studied species at diferent alu’, Scie Altitude urep HTCP Py opttemp) Py max) Eschalic 2950 m $1.1 3052216 $000 9508 781 7981.0 H taicoider 2950 m “sors = 330416 S24 400m = 25808 220s ass 3550 m S403 3103413 4200 m “easio 333816 A seosus 3550 m 204 alg +4200 2sH18 soso 1.7805 ALTITUDE (m) Fc 3. - Optima lemperature fr photasynesis at iferent aitudes for species in bth scan peta schlei (@), Hypericum aricoides (C), Calandrinia acculis (8) and ypochoeis toes (A) The high temperature compensation points also show interesting results. At 4000 m, this temperature for the high level plants is 22°C for H. laricoides and 25°C for E. schultzii. While at 4-200 m the two acaulescent rosettes show compensation points at much higher temperatures, 31°C and 33°C for H. setosus and C. acaulis, respectively (figures 1 and 2, table 1). With respect to low temperature compensation points there are no clear trends between strata, E. schultzii shows low temperature compensation point of approximately — 10°C for both altitudes while the other three species have much higher points, H. sefosus has the highest points with approximately ~3.0°C for the two altitudes, The differences in optimum temperature for photosynthesis and altitude for the chosen species are therefore well differentiated if we compare the different strata (fg. 3). The two higher strata species show an expected decrease in leaf temperature optimum for photosynthesis with increase in altitude. Surprisingly enough, one of the acaulescent species, H. setosus shows an opposite trend, while the other, C. acaulis maintains similar values along the gradient. Discusston Many authors have described how leaves of alpine plants reach impressively high temperatures compared to air temperatures. Gares & Janke (1966) describe leaf temperatures of 8 to 10°C above air temperature for Polygonum bistortoides i the Rocky mountains. Sauisaury & Spon (1964) find leat temperatures 10 to 15°C WL IB, n* 5 - 1992540 . Rada et al above air temperature for different Rocky mountain plants at 3 $00 m. Koawen & Cocraxe (1983) working with different life forms at 2 040 m in the Snowy Mountains of Australia find that the smallest differences between daytime leaf and air temperatures occur in trees and shrubs (between 0 and 10°C); sessile rosettes have mean differences of 14.5°C and maximum differences of up to 24°C and cushion plants have the greatest differences with a mean of 27.4°C and maximum of 30°C. These last authors suggest that these differences nearly compensate for the altitudinal drop of maximum air temperature. Although we did not measure field leaf-air temperature differences in this study, we have found for other species quite similar results as those of Koeex and Cocirane (1983). These differences range from 0 to 10°C in Espeletia spicata and Polylepis sericea, a giant rosette and a small tree, respectively (Rapa, 1983); from to 15 to 25°C in Hinterhubera lanuginosa and ‘Arenaria jahnii, a dwarf shrub and a cushion plant, respectively (unpublished data); all of them growing at 4 200 m in the Paramo de Piedras Blancas. We may then suppose that the plants chosen for this study behave in a similar manner in terms. of daytime leaf temperature. We now may ask ourselves: how do these leaf-air temperature differences influence on the process of photosynthesis for each of these life-forms? Our results show how each of the species adapts to the microenvironment where it is found. Having in mind that E. schulteii and H. laricoides grow at 50 to 120 cm above ground level, the lowering of the leaf temperature optimum for photosynthesis is coupled to the lowering in air temperature along the gradient. On the other hand, Hz setosus and C. acaulis, growing between 0 and 2 cm are greatly influenced by soil temperature and as a consequence optimum temperature for photosynthesis is similar or it even increases with altitude since bare soil at the higher altitude warms up much more than atthe other altitude. Similar results have been described for other species in other regions. Siarvex (1978) shows that for Eucalyptus paucifiora, in the Australian Snowy Mountains, the optimum temperature for photosynthesis follows the normal air temperature decrease while other authors have found different results, for low growing plants. Koawex & Drewex (1987) find similar optimum temperature for photosynthesis in different herbaceous plants from the Alps along an altitudinal gradient, With respect to low limits for net photosynthesis, it is interesting to note that all four species show positive assimilation rates at below 0°C forall altitudes. Together with these four species, Lucilia venezuelensis, a cushion plant, growing at 4 200 m showed positive net photosynthesis at temperatures of ~5 to ~7°C (unpublished data). Other authors (Pisrx et al., 1967; Tsszex et al., 1981; taken from Korwe & Laxcuse, 1988) obtain CO, gains in high alpine forbs down to -5°C. Espeletia schultzii, a'freezing intolerant species (RaDs et al., 1987) shows carbon gain down to approximately ~10°C. This supports Korxer & Larcis's (1988) observation on net photosynthetic CO; uptake being very close to the low temperature limit of leaf survival in species which avoid freezing. Cold resistance mechanisms would have to be studied for the other three species. In the case of high temperature limits for photosynthesis, our results are below those found in the literature. We ‘obtain carbon gains at temperatures of up to 30-34°C for all species, contrasting those results of Lancue and Waovsr (1976) who report values from 38 to 47°C for different species; including 42°C for Espeletia semiglobulata compared to 31°C at 2.950 m and 25°C at 4 000 m for Espeletia schultzii in our study. Acta Eeotgica [Net photosynthesis ~ Leaf temperatur A final consideration would rates. At optimum temperature show higher assimilation rates (3-5 mol ms") in this study et al., 1989). A possible expl from’ soil respiration for the g are required to confirm this hy We may conclude this wor of the temperature dependene dependent on the type of life and more important “areas typi “cold” for all plants and their r ACKNOWLEDGEMENTS J. Gowzatez" stay inthe CTELAT Latinoamercana de Botanica fellowshi ‘rant (CDCHT-ULA C-444-90), REFERENCES ‘Azocan A. & MoxAsteno M., 1980, — de Macubaji. I: Monasrixio M., los Andes, Merida, 285-252 Azocas A, RADA F. & GoLDstEN G., ‘aulescentrosete species, Orcolg (Cor M. 1, 1967, ~ The ecology ofthe a Cusmercadas J, 1979. Growth forms high tropical Andes. fm Lansow K Press, 397-410 Pin C. Bi, BALL J.T. & BERRY J. A 1 RW. Ensnisoer I, Moosey H ethos and instrumentation. Chap GGares D.M. & Tanke R, 1966. ~ Thee Garnernis G., Raoa F Azoeak A. 19 in andean giant rosete species. Oe Gotpsrens G., RADA F., CANALES M. & aulescontroseue species, col Henge 0, 1964, ~ Features of foal Kowa C. & Cocimans P.M, 1983, — in summer along an elevational gr Komsa C. & DINER M,, 1987, = Insta in herbaceous plants from low and Konan C. de Lance W., 1988, ~ Pla Plants and Temperate. Symp. 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A possible explanation could be from soil respiration for the ground level plants ‘te required to confirm this hypothesis. iWe may conclude this work quoting Konwen & Laxcnen (198 of the temperature dependence of éEpendent on the type of life process (daytime or nightime activin and more important “areas typically classified as cold climates ane ‘cold’ for all plants and their respective functions” spect to maximum photosynthetic 8, the two acaulescent rosettes ACKNOWLEDGEMENTS: 3. GowzaLer stay in the CIELAT CU atinoamericana de Botanica fellowship. punt (CDCHT-ULA C-444-90), nivesidad de los Andes, Venezuela) was supported by a Red ‘This research was supported by a Universidad de los Andes REFERENCES Atocan A. & Mowastenio M, 1980, ~ de Mucubaf. re Movastenio M, les Andes, Merida, 255-262, ‘Goel Rana F. & Coupee G., 1988. ~ Freezing tolerance in Draha chonopila, a “miniature” aulescent rosete species. Oecologia, 78, 156-160; Cee 97. The ecology of the alpine zone of Mt. 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Soito F, RaDA F.& Azocax A, 1991, Freezing tolerance and avoidance in high wopcalandean plans sit eqully represented in species with different plant height!” Oecologia, 86, 378-382 ‘Tse L. L, Liwns MC, MILER. C, Ma¥0 I, CHAPIC ITF. S, & Osx. W., 1981. ~ An analysis of process of primary proditon in tndra growth forms. /m: Buss L.C.. CRoca J. B.. Heat D. W. & Moone 5 J. tds. A Comparative Analysts of Tundra Econytems.Iteroatonal Biokogical Progranune, 25, Cambridge: Cambridge University Press, 285-356, Acta Ecologica ‘Acta Ecologica, 1992, 13 (5), 543-552. The within-panick Laborato Goure ORS Abstract The total umber of spkelt Aistebation slong the panicle axis, Within or between clones, The see sizeof the panicles of the clone. day, Is relation to panicle size is energy are also quadratic. The mu is discussed in relation to culm be ‘wopical envzonment, Keywords: Panicum maximum, ps Résumé Le nombre total piles d's cence Laréparition des Eilts de tale pees, entre petites et grand Propagation bape do la frase ea panicule Le nombre ¢ eile ‘uadratigue de a ale de infor rméme ordre. La rupli ds st en relation ave apparition succes sux fluctuations de pluvionére e INTRODUCTION Panicum maximum is an é Tanzania (Comes, 1975; Pent tropical regions, its commercializ (Husenneys, 1975). As with many and shedding at maturity constt Acta Ccologica 1146.69 XRIOSISSVK