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Quercus Petraea and Q. Robur) to Moderate Drought Frank Thomas, Thomas Gausling

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Quercus Petraea and Q. Robur) to Moderate Drought Frank Thomas, Thomas Gausling Morphological and physiological responses of oak seedlings (Quercus petraea and Q. robur) to moderate drought Frank Thomas, Thomas Gausling To cite this version: Frank Thomas, Thomas Gausling. Morphological and physiological responses of oak seedlings (Quer- cus petraea and Q. robur) to moderate drought. Annals of Forest Science, Springer Nature (since 2011)/EDP Science (until 2010), 2000, 57 (4), pp.325-333. 10.1051/forest:2000123. hal-00883316 HAL Id: hal-00883316 https://hal.archives-ouvertes.fr/hal-00883316 Submitted on 1 Jan 2000 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Ann. For. Sci. 57 (2000) 325–333 325 © INRA, EDP Sciences Original article Morphological and physiological responses of oak seedlings (Quercus petraea and Q. robur) to moderate drought Frank M. Thomas* and Thomas Gausling Universität Göttingen, Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Abt. Ökologie und Ökosystemforschung, Untere Karspüle 2, D-37073 Göttingen, Germany (Received 13 September 1999; accepted 4 January 2000) Abstract – To study their responses to a moderate drought, seedlings of sessile (Quercus petraea [Matt.] Liebl.) and pedunculate oak (Q. robur L.) were grown in outdoor lysimeters that allowed unrestricted root growth. They were subjected to a moderate drought from mid-June to the end of August. The stomatal conductance remained almost unaffected by drought. In the drought-stressed seedlings, the relative foliar water deficits were significantly higher, and the predawn leaf water potentials were significantly lower than in the control plants. The decrease in osmotic pressures at maximum turgor and at the point of turgor loss in the drought-stressed plants indicated osmotic adjustment. The ratio of leaf to fine-root biomass was significantly diminished in the drought-stressed seedlings. In combination with previous observations, the results indicate that, in the Central-European oak species, a shift in biomass compart- mentation is the predominant process in acclimation to drought since it also occurs under severe drought stress in contrast to osmotic adjustment, which was only found under conditions of a moderate drought. Quercus / drought / leaf-root biomass ratio / osmotic adjustment / pressure-volume analysis Résumé – Réponses morphologiques et physiologiques de semis de chêne (Quercus petraea et Q. robur) face à une sécheresse modérée. Afin d’étudier leur réaction face à une sécheresse modérée, des plantules de chêne sessile (Quercus petraea [Matt.] Liebl.) et de chêne pédonculé (Q. robur L.) ont été cultivées à l’extérieur, dans une cuve lysimétrique permettant un développement illimité des racines. Le régime de sécheresse modérée leur a été appliqué de mi-juin jusqu’à fin août. La conductance foliaire stomatique n’a guère été affectée par la sécheresse. Le degré du déficit hydrique foliaire relatif des plantules stressées par la sécheresse avait aug- menté de façon significative alors que le potentiel hydrique foliaire mesuré avant l’aube fut plus bas que celui des plantules témoins. La diminution des pressions osmotiques en pleine turgescence et au point zéro turgor des plantules stressées indiquerait un ajustement osmotique. Les plantules stressées ont montré un ratio de la biomasse foliaire par rapport à la biomasse des racines fines qui avait baissé de manière significative. Les résultats complètent ainsi des études antérieures indiquant que pour les chênes européens le changement de la compartimentation de la biomasse serait le processus prédominant dans l’acclimatation à la sécheresse. En effet, ce phénomène surgit également lors d’une sécheresse excessive tandis que l’ajustement osmotique n’a été constaté que lors d’une séche- resse modérée. Quercus / ajustement osmotique / analyse pression-volume / rapport des biomasses feuilles-racines / sécheresse * Correspondence and reprints Tel. +49-(0)551-395724; Fax. +49-(0)551-395701; e-mail: [email protected] 326 F.M. Thomas and T. Gausling 1. INTRODUCTION a slower rate of soil dehydration. This indicates that the degree of the stress affects the performance of osmotic Besides beech (Fagus sylvatica L.), pedunculate adjustment [5]. However, in that particular study, the ∏ (Quercus robur L.) and sessile oak (Q. petraea [Matt.] osmotic pressure at maximum turgor ( o) was measured Liebl.) are the most common deciduous forest tree in sap which had been pressed out of the leaf tissue. Due species in Central Europe. Within this region, under nat- to dilution of the cell sap by apoplasmic water, this ∏ ural conditions, they have a lower competitive power than method generally leads to an overestimation of o (i.e., beech at sites with a moderate to optimum supply of less negative values [20]). Seedlings of the pedunculate water and nutrients. Therefore, they tend to gain domi- oak were shown to osmotically adjust their leaf water nance over beech mainly at sites with more extreme soil potential when subjected to consecutive drying cycles moisture conditions [8]. Although, compared to the ses- [19]. However, these plants were grown under green- sile oak, the pedunculate oak was found to have a larger house conditions, and in containers with restricted rooting water requirement [17] and to be more sensitive to volume as was also the case in other studies [9, 28]. drought stress [4, 27], this species also occurs at sites with Therefore, we conducted a study on pedunculate and ses- low water supply, e.g. on nutrient-poor sandy soils sile oak seedlings whose roots were allowed to grow (Betulo-Quercetum roboris; [29]), or on plateaus and without restriction outside the greenhouse under con- exposed slopes of well-drained limestone hills (xerother- trolled conditions. The objectives of the investigation mic mixed oak forests; cf. [13]). At those sites, drought were to examine whether (1) osmotic adjustment would tolerance is a prerequisite for the establishment of develop under a moderate drought, and (2) morphological seedlings. alterations as responses to a moderate drought would be similar to those caused by severe stress. The results are Morpho-anatomical and physiological mechanisms of discussed against the background of findings from a pre- drought tolerance are known. Morpho-anatomical mecha- vious investigation on the effects of a severe drought on nisms involve alterations in leaf area, cell wall elasticity, oak seedlings [24]. or biomass compartmentation between shoot and roots [15]. Physiological mechanisms include an effective stom- atal control of water loss and osmotic adjustment, i.e. a 2. MATERIALS AND METHODS change in osmotic pressure larger than the passive change resulting from the loss of cell water [11]. In Q. petraea, 2.1. Plant material and cultivation severe drought resulted in an increased production of fine- root biomass, and in a decrease in the following: the bio- For the experiments, selected reproduction material mass of leaves and stems, the biomass ratio of leaves to originating from the north German heathland and hill fine roots, and the number of buds. In Q. pubescens, with country (sessile oak (Quercus petraea [Matt.] Liebl.); regard to biomass parameters, only the average mass of derived from the Forest Seed Centre in Oerrel, Lower the individual buds was significantly lowered by drought; Saxony), or originating from the hilly and mountainous but, in this species, indications were found for an increase country of southern Germany (pedunculate oak (Quercus in tissue elasticity as an acclimation to drought [24]. robur L.); obtained from a tree nursery) was used. In However, in neither of the two species was osmotic February 1998, the acorns were sawn into germination adjustment found as a response to severe drought [24], nor dishes filled with quartz sand and cultivated in a green- was it detected in most of the drought-resistant shrubs, house with a temperature of 15 °C and a photosynthetic including Q. pubescens, that were investigated at drought- photon flux density (PPFD; Osram HQJE, 400 W) of 135 prone sites in Austria and the Mediterranean region [11]. to 210 µmol m–2 s–1 (day; 6.00 – 20.00 h), 10 °C (night), In contrast, indications for osmotic adjustment as a means and a relative humidity of 50 to 85%. In mid-May, when of acclimation to drought was provided by a study carried the shoots were approximately 10 – 15 cm tall, the out in a mixed stand of F. sylvatica, Q. petraea and seedlings were planted into four lysimeters situated in the Q. pyrenaica in central Spain [3]. Botanical Garden. These were 5 m long, 1.2 m wide and Osmotic adjustment may occur during ontogeny, and 1–2m deep, and were filled with calcareous sand not only in response to drought [2]. Therefore, the (pH(H2O) approx. 7). In each lysimeter, 60 seedlings assumption of osmotic adjustment needs to be corrobo- (30 sessile and 30 pedunculate oaks) were arranged in rated by comparisons between drought-stressed and four rows at a distance of approximately 25 cm from the adequately watered plants, which are in a comparable adjacent plants or from the side of the lysimeter to pre- developmental stage. In a recent study conducted on vent shading effects. The oaks were thoroughly watered, seedlings of sessile oak, osmotic adjustment was less pro- and the water level in the lysimeters was adjusted to nounced after a rapid onset of drought than in the case of 40 cm below the soil surface. During the first two weeks Responses of oak seedlings to moderate drought 327 after planting, the seedlings were protected from inten- To supply the plants with nutrients, 10 l of nutrient sive solar radiation by shading.
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