Báo cáo khoa học: "Influence of soil drying on leaf water potential, photosynthesis, stomatal conductance and growth in two black pine varieties"

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Original article drying on leaf water potential, Influence of soil photosynthesis, stomatal conductance and growth in two black pine varieties Gérard Lévy Gilbert Aussenac b , c François Lebourgeois a Bernard Clerc François Willm c , c Équipe a phytoécologie, Unité d’écophysiologie forestière, Inra Nancy, Champenoux, France 54280 b sol et nutrition, Unité d’écophysiologie forestière, Inra Nancy, Équipe 54280 Champenoux, France Unité d’écophysiologie forestière, Inra c Équipe bioclimatologie, Nancy, 54280 Champenoux, France accepted 20 October 1997) 12 June 1997; (Received Abstract - The aim of this study was to examine the influence of long-term soil water deficit on growth and physiological processes of two black pine varieties (Pinus nigra ssp. laricio var. ssp. laricio var. Calabrica). Three-year-old seedlings grown in larges Corsicana and Pinus nigra ) 3 m were subjected to a prolonged summer drought (99 days from the end boxes (volume: 1.62 of June until the end of September) and photosynthesis (A), stomatal conductance (g water sta- ), w tus and growth were measured. No marked differences arose between Corsican and Calabrican pines feature to drought. At least in their juvenile stage, both varieties exhibited a ’drought- avoidance strategy’ characterized by an efficient stomatal control of transpirational water loss. This result is consistent with previous studies on Pinus nigra and confirm the water stress adapta- tion of this collective Mediterranean species. Because a significant decrease of g (about 30 %) w the data suggested that predawn water potential was observed with no obvious variation in , wp Ψ was not the best indicator to precociously detect water stress. However, both A and g reduced w to nearly zero as soon as the threshold value of Ψ = -1.6 MPa was reached (respective values wp 0.5 μmol m s and 11 mmol m s Because most fine roots were within the upper 40 cm of -2 -1 -2 -1). the soil, a superficial soil desiccation has probably induced rapid stomatal closure, triggered by a biophysical and/or biochemical signal from the desiccated roots to the leaves. Embolism seems not to be responsible for the effect of drought on physiological processes, because the minimum observed at the end of the drying cycle (-2.5 MPa) remained higher than the thresh- value wp of Ψ inducing a significant xylem cavitation for these varieties (-4 MPa). Summer drought sig- old nificantly reduced annual stem diameter (-20 %) and needle length (-25 %), but not stem elon- gation. Total elaborated dry weight was reduced about 45 %. Seedlings grown in the dry regime * Correspendence and reprints E-mail: lebourgeois@nancy.inra.fr
reduced belowground growth proportionally more than aboveground growth, causing a signifi- cant decrease in the R/S ratio. Such a result, which diverges with classical models of whole plant biomass partitioning, might be partially explained by seasonal pattern in the root growth which typically has its most important peak in mid-summer, period of maximum drought in our study. With the parameters studied here, the expression of the genetic characteristics between varieties in drought tolerance appeared to be limited. Thus, further investigations could be undertaken to learn about drought feature at cell and molecular levels. (© Inra/Elsevier, Paris.) Corsican pine / Calabrican pine / leaf conductance / photosynthesis / drought / root/shoot / growth / biomass partitioning /Pinus nigra Résumé - Influence de la sécheresse sur le potentiel hydrique foliaire, la photosynthèse, la conductance stomatique et la croissance de deux variétés de pins noirs. L’objectif de cette étude était d’analyser l’influence d’une sécheresse édaphique prolongée sur la croissance et le com- portement écophysiologique de deux variétés de pins noirs (Pinus nigra ssp. laricio var. Corsi- cana et Pinus nigra ssp. loaricio vsar. Calabrica). Des plants99 ej 3(dens, cjuin à findans des grandes d ultivés a ), 3 m nt été oumis à une sécheresse de cuves (volume : 1,62 fin septembre) pen- dant laquelle la photosynthèse, la conductance stomatique, l’état hydrique et la croissance des plants ont été mesurés. Au moins dans leur stade juvénile, les deux variétés ont présenté la même stra- tégie « d’évitement» caractérisée par un contrôle stomatique efficace de la transpiration. Ce résultat est conforme à ceux obtenus sur les pins noirs et confirme la stratégie commune d’adap- tation à la sécheresse de cette espèce méditerranéenne. La fermeture stomatique rapide, avant que le statut hydrique ne soit affecté, suggère que le potentiel hydrique de base n’est pas le meilleur paramètre pour détecter précocement le stress. Cependant, la photosynthèse et la conductance sto- matique se sont stabilisées à des valeurs très faibles dès que le potentiel hydrique de base a atteint le seuil de -1,6 MPa (respectivement 0,5 μmol m s et 11 mmol m s Le système -2-1 ). -2 -1 racinaire superficiel (densité maximale de racines dans les 40 premiers centimètres) a probable- ment joué un rôle déterminant en détectant précocement la sécheresse puis en transmettant un signal chimique et/ou biophysique des racines sèches jusqu’aux feuilles. La cavitation ne semble pas avoir joué un rôle majeur dans les comportements observés étant donné que la plus faible valeur mesu- rée de Ψ (-2.5 MPa) est restée supérieure au seuil d’embolie défini pour cette espèce wp (-4 MPa). L’accroissement radial et la longueur des aiguilles ont été significativement réduits par la sécheresse estivale (respectivement de -20 et de -25 %) alors que l’élongation de la pousse ter- minale n’a pas été affectée. La réduction totale de matière sèche élaborée a été d’environ 45 %. Les plants soumis à la sécheresse ont alloué moins de ressources à la croissance racinaire qu’à la croissance aérienne induisant une diminution significative du rapport R/S. Un tel résultat diverge des modèles classiques d’allocations de matières, mais peut partiellement s’expliquer par le rythme saisonnier de croissance des racines qui atteint son optimum au milieu de l’été ; période de sécheresse édaphique maximale dans notre étude. Avec les paramètres étudiés ici, l’expression des variations génétiques entre les deux variétés dans le comportement vis-à-vis de la séche- resse est apparue limitée. Ainsi, d’autres travaux devraient être envisagés afin de mieux cerner les régulations aux niveaux cellulaire et moléculaire. (© Inra/Elsevier, Paris.) laricio de Corse / pin laricio de Calabre / conductance stomatique / pin photosynthèse / sécheresse / R:S / croissance / répartition de biomasse /Pinus nigra 1. INTRODUCTION their natural stands, the ecological growth conditions are fairly similar with a wide range of altitudinal and soil conditions and Calabrican pine (Pinus nigra ssp. lari- a hot, dry summer [ 16, 17, 19]. The mor- cio var. Calabrica) and Corsican pine phological differences between these vari- (Pinus nigra ssp. laricio var. Corsica) are two varieties of the Mediterranean col- eties are small, particularly among mature lective species Pinus nigra [10, 50]. In trees, and apply to bark structure, leaf mor-
highlight the underlying mechanisms to phology and anatomy, as well as cone morphology [3, 21, 22]. Despite the the growth decrease observed in mature increased interest in their use for afforesta- stands, an ecophysiological study was tion, their ecological plasticity outside the undertaken. In this present work, plant area where they are indigenous is not well water status, gas-exchange responses and known. In France, Corsican pine is gen- growth of both irrigated and droughted erally recommended on more or less acidic seedlings were measured and analysed. soils, whereas Calabrican pine seems to be able to withstand soils with temporary water table [23, 36]. However, both vari- eties are mixed in most managed crops. 2. MATERIALS AND METHODS Pinus nigra water stress tolerance is also poorly understood and comparative 2.1. Experimental design studies between subspecies and varieties of this species are rather rare. Becker [7], The experimental design was set up at the studying transpiration and drought Inra Research Centre of Nancy (Lorraine, behaviour of 3-year-old seedlings of some northeast France). It consisted of four large coniferous species (Eastern White pine, partially buried boxes (depth: 100 cm; width: Douglas fir, Norway spruce and Corsican 144 cm; volume: 1.62 m Thus, the root ). 3 pine), showed that Corsican pine had the development of plants was not limited or at best water use efficiency (shoot biomass the least less confined than if they had been in increment versus transpired water during small-sized containers. These boxes were filled the experiment). Aussenac and Granier with 10 cm of gravel at the bottom to improve water drainage, and 80 cm of a sifted sandy [5] and Aussenac and Valette [6] showed clay loam soil from the horizon A1/A2 of a that in response to soil water depletion, Dystric Cambisol (Food and Agriculture Orga- 15-year-old black pine trees (Pinus nigra nization classification) from the Haye Forest ssp. nigricans) rapidly decreased transpi- (France). The characteristics of the growing ration and photosynthesis, with these pro- substrate are presented in table I. No fertiliza- cesses being totally inhibited at a rela- tion was applied because chemical composi- tively high predawn needle water potential tion corresponded to optimal conditions of (-1.6 to -1.7 MPa). For the subspecies plant nutrition [36]. ’laricio’, Aussenac [4], using excised shoots subjected to a desiccation, beginning of March 1992, 2-year- At the described a similar feature with a signifi- old (2 0) seedlings (seed origin: Corsica and + Calabrica; average height: 10 cm) from the cant stomatal closure when predawn nee- Forest Research Center’s nursery were planted dle water potential reached values around in staggered rows (35 plants of each variety -1.2 to -1.4 MPa. A dendroecological per box). In order to avoid any possible inter- study carried out on 1 808 mature Corsican variety competition effects, varieties were not pine in western France also showed that mixed in the boxes. During this first growing drought was one of the major environ- season ( 1992), all trees were grown in open mental factors influencing and limiting conditions, and kept well-watered by natural radial growth and wood productivity. The irrigation. No herbicidal and manual fertil- or current decline of trees has been mainly izing treatment was applied. related to repeated severe drought events that have occurred since the end-1960s in At the beginning of June of the following year (1993), a transparent polyethylene tunnel this region [35]. opened at its extremeties was installed in order In order to compare the water stress to intercept rainfall and to maintain sufficient sensitivity of both varieties and also to ventilation during hot summer days.
2.2. Water supply mental conditions during the season were: regimes 1054 ± 436 μmol m s -2 -1 air ;T 25.2 PPFD = = (± 2.8) °C; air CO concentration (C 346.6 2 ) a = The consisted of two experimental plots (± 14.5) μmol mol Net CO assimilation rate . -1 2 control plots and two dry plots. (A, μmol m s and stomatal conductance -2 -1) Irrigated plots: maintained permanently near to water vapour (g mmol m s were cal- -2 -1) , w field capacity by frequent manual watering. culated with the classical equations of Caem- Soil water content was measured weekly in merer and Farquhar [9]. Gas exchange mea- each box at a depth of 40 and 60 cm with two surements were made at the same time as tensiometers. midday leaf water potential. At the end of the experiment, calculations were performed on Droughted plots: to extrapolate to natural the basis of the total projected needle area of the conditions, a prolonged summer drought was branches using a video camera coupled to an imposed. Drought began on 22 June 1993 image analyser (ΔT Devices, Cambridge, UK). (Julian day 174). The rewatering to field capac- ity occurred on 2 October (Julian day 276), after 99 days of drought. Seedling recovery was sampled 3, 6 and 10 days after rewater- 2.4. Growth measurements ing. Soil water content was measured weekly in each box at a depth of 40 and 60 cm with two psychrometers. Unfortunately, due to technical At the end of the first growing season (Octo- problems, soil water potential could only be ber 1992), annual stem elongation and total measured from day 36 of the drying cycle. height were measured. In 1993, the bud expansion was first observed in early April. The first stem mea- 2.3. Ecophysiological measurements surement was made in mid-June before the beginning of the moisture stress treatment. After the summer drought, total height, total Ecophysiological measurements were car- stem elongation, basal stem diameter and length ried out on 16 3-year-old seedlings (four plants of new needles were measured. The needle of each variety per treatment) representative length was measured to the nearest mm from of the sample. The leaf water potential was the point of fascicle sheath insertion in the axil measured weekly from 22 June onwards on of a subtending cataphyll (bract) to the needle needles using a pressure chamber [46]. Nee- top. For each plant, ten needles were randomly dles were sampled in the middle of the annual chosen in the middle of the current-year shoot. shoot just prior to dawn (predawn leaf water potential, Ψ and at 1 pm solar time when ) wp In order to estimate the biomass distribu- the sun was at its zenith (midday leaf water tion in the various organs (needle, stem and potential, Ψ ). wm root) during an annual vegetative cycle and to Gas exchange measurements were per- quantify the below- and aboveground growth formed using a portable gas exchange mea- responses to the water stress treatment, two surement system (LiCor 6200, LiCor, Lincoln, boxes (one control plot and one dry plot) were NE, USA) under natural climate. Environ- harvested at the end of the drought. Because
of the absence of blocking (box) effects, the 3. RESULTS two boxes were randomly chosen. For the root system, each plant was manually and carefully 3.1. Plant water status uprooted, and to avoid any error due to wall effect, plants near the wall were eliminated. No marked differences arose between However carefully applied, this method did Corsican and Calabrican pine leaf water not allow us to sample all fine roots. Never- drought (figure 1). potential response to theless, direct and visual observations showed From 15 days onwards, water stress that the root system remained superficial (max- increased gradually with predawn water imum root density above 40 cm) for both treat- ments and varieties. Only the largest roots potentials decreasing around -0.2 MPa reached the deep soil horizon (1 m). All the per week in both varieties. After 73 days, samples were also oven-dried at 80 °C for 48 h. predawn water potentials reached stable The biomass partitioning among the plant com- values around -2.5 MPa. The decrease in partments was assessed by determining a) the Ψ was closely related to soil wp water con- leaf mass ratio (LMR, leaf dry mass/whole tent (figure 2). plant dry mass, g g b) the stem mass ratio ), -1 (SMR, stem dry mass/whole plant dry mass, In watered treatments, and Ψ wm wp Ψ g g c) the root mass ratio (RMR, root dry ), -1 from -0.22 to -0.55 MPa (aver- ranged mass/whole plant dry mass, g g d) the ), -1 age: -0.32 ± 0.08 MPa) and from -0.7 to root/shoot ratio (R/S, root dry mass/shoot dry -2.2 MPa (average: -1.2 ± 0.4 MPa), mass, g g The shoot dry weight equalled ). -1 respectively. These values corresponded to the dry weights of leaves plus stems plus branches. the common observed data for the Pinus species [44]. One-way and two-way analyses of variance After rewatering (R), seedlings recov- (ANOVA followed by Fisher’s PLSD test) ered rapidly. Three days after rewatering, were used to evaluate the significance of the water potentials were again equivalent to single and interactive effects of drought and those of irrigated plants (figure 1). varieties.
3.2. Stomatal conductance and net and A reached values wp when Ψ near zero CO assimilation rate reached -1.6 MPa. 2 As illustrated in figure 3, under water 3.3. Plant growth and dry matter stress stomatal conductance (g decreased ) w rapidly in both varieties. The decrease in In 1992, no mortality or visible dam- photosynthesis (A) occurred later when age was observed at the end of the first g presented a decrease of about 50 % of w growing season. However, the first sea- the initial values (below 30 mmol m s -2 -1 ) son terminal shoot growth was signifi- (figure 4). For both varieties, g and A w cantly lower for Calabrican pine than for stabilized around minimal values of Corsican pine (table II). 11mmol m s and 0.5 μmol ms -2 -1 -2-1 after 43 days of drought. Rewatering In both varieties, water stress during induced a rapid recovery of stomatal con- the second growing season had no influ- ductance and CO assimilation. 2 ence on annual stem elongation, but reduced significantly stem diameter (mean In watered treatments, g and A showed w value for both varieties: -20 %) and length considerable variability due to plant-to- of new needles (mean value for both vari- plant variability and weather fluctuation eties: -25 %) (table III). Under well- during the season. Through the season, watered conditions, a significant differ- the average values were 76.9 mmol m -2 ence was also noted on needle length -1 s and 2.9 μmol m s for Calabrican -2 -1 which appeared shorter in Calabrican pine pine and 79.0 mmol ms and 3.6 μmol -2-1 than in Corsican pine (-12 %) (table III). -2 -1 m s for Corsican pine for g and A, w That was expected because needle length respectively (figure 4). is one of the morphological differences between these two varieties [18]. Both varieties showed a similar evolu- tion of A and g in response to decreas- w In both varieties, total elaborated dry (figure 5). Stomatal conductance ingwp Ψ in the dry regime averaged less weight decreased sharply between -0.4 and -1.1 than 60 g, while elaborated biomass in the MPa. Inhibition of A started below -1.1 irrigated plots averaged over 95 g (table MPa but dropped rapidly thereafter. g w IV). Drought was responsible for a
decrease of about 51, 36 and 42 % in stem, Mediterranean habitats characterized are needle and root elaborated dry matter and ecological by great climatological diversities which have induced differential respectively. drought adaptations in the different species marked decrease in Drought led to a [40]. Thus, a lot of other Mediterranean SMR, RMR and R/S ratio and to an species exhibit a drought-tolerance strat- increase in LMR (table IV). Moreover, in egy: Cedrus atlantica, Quercus pubescens. both treatments, RMR was lower in Cor- Quercus ilex, Buxus sempervirens, etc. sican pine than in Calabrican pine. At the [20]. Thus, it appears that there is no com- end of the drying cycle, no massive leaf mon Mediterranean strategy to moisture abscission was observed. stress response. As previously observed on Norway 4. DISCUSSION AND spruce [11, 37] and in several oak species CONCLUSIONS [8, 42], care must be taken with the use of predawn water potential as a driving Three-year-old Calabrican and Corsican variable of stomatal closure. In our study, pines exhibited similar decrease of net a large decrease of g (about 30 %) was w CO assimilation rate and stomatal con- observed with no obvious variation in 2 ductance with increasing drought. Our = -0.4 MPa) predawn water potential wp (Ψ results are in agreement with previous that this parameter is not suggesting work on Pinus nigra [4-7, 32] and con- always the best precocious indicator of firm the water stress adaptation of this col- the water stress actually experienced by lective Mediterranean species which plants. However, when drought increases, appears to be linked to the ability to avoid the close correspondence of predawn with internal water stress. Similar features have soil water potentials supports their use as been obtained in a wide range of Pinus an indicator of soil moisture. There is still and Mediterranean species: Pinus unci- considerable uncertainity over the under- lying causes of drought-induced changes nata and Pinus pinaster [6], Pinus sylvestris [45], Abies bornmuelleriana in plant feature. However, in our study, we may suggest that the superficial root [25], Quercus afares and Quercus faginea system of the seedlings has probably [1]. However, it must be noticed that the
played an important role in the early vari- applied during the maximum stress was growth period, which could explain ations of g by precociously detecting the w root increase in soil hydraulic resistance [14, why the dry regime reduced belowground 39, 47].Many recent studies ascribe stom- growth proportionally more than above- atal closure to a single chemical substance, ground growth, causing a significant such as cytokinin or abscisic acid, trans- decrease in the R/S ratio. This response ported from the roots in the drying zones diverges with models of whole plant of the soil to the leaves by means of xylem biomass partitioning that predict that low rates of water absorption will result in flux [13, 15, 49]. In our study, decrease in plant hydraulic conductance through increased biomass partitioning to root growth [ 12, 28, 38, 41]. However, it is cavitation and embolism in the xylem con- duits [30, 48] seems not to be responsible clear that partitioning of assimilates between roots and aerial parts may differ for the effect of drought on physiological processes, because the minimum value of under long- and short-term drought and that water stress also has a greater effect predawn potential (-2.5 MPa) water remained higher than the threshold of during certain phases of the plant’s cycle -4 MPa inducing a significant xylem cav- than others [31]. For the root system, although the differences were small and itation for these varieties [24]. not always statistically different (at After the first growing season, height P < 0.05) (table IV), the data might suggest growth differences suggest a more severe a greater investment in root growth in Cal- transplanting shock for Calabrican pine. abrican pine than in Corsican pine. From The mechanisms underlying these differ- the perspective of morphological adapta- remain unclear but may involve root ences tion to drought, further investigations regeneration and elongation decrease over could be undertaken to confirm this trend. the first planting year [26]. restored stomatal Rewatering rapidly As expected, the summer water stress conductance, photosynthesis and water had a significant impact on growth and well-watered conditions. In addi- status to severely reduced whole-plant biomass tion to a capacity for avoidance of dehy- accumulation by a factor of 1.9 in both dration through stomatal closure, the rapid varieties. The annual stem elongation was recovery in gas exchange might also indi- not affected by drought because most of a robust, dehydration-tolerant photo- cate the elongation occurred during the spring synthetic apparatus. Nevertheless, a sig- when water was available in large nificant shoot growth decrease was Terminal buds started to break in amounts. observed the following year [34], sug- and stopped growing by late early April gesting important after effects of whereas radial increment continued June, unfavourable climatic conditions tree on till September [34]. This phase of growth growth. is in agreement with previous results and confirms the monocyclic shoot growth in In conclusion, the pine plants were Pinus nigra [27, 29, 33]. The seasonal pat- characterized by a high sensitivity to tern in root growth has not been investi- drought associated with an efficient stom- gated in the present study. However, Cor- atal control of transpirational water loss. sican pine typically has two peaks in root With the classical ecophysiological param- growth. A low activity during spring till eters studied, the expression of the genetic characteristics between varieties in drought the end of June followed by a period of important root growth from July to tolerance appears to be limited. This pre- liminary study suggests further investiga- September [2, 43]. Based on these obser- vations, we may suggest that the water tions which could be undertaken to learn
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