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and thus leaf?and 57 even whole tree?functionality during drought, is related to 58 stomatal aperture Leaf stomatal conductance is driven by 59 atmospheric water pressure deficit and soil water supply. 60 By closing their stomata during drought, trees reduce leaf 61 transpiration and therefore water-use, but in the mean time, 62 CO 2 uptake and assimilation rates are restricted. 63 Differences in structural characteristics among tree 64 species contribute to the considerable variability in their 65 responses to water depletion in drought-exposed forests. 66 Among these features, the vertical development of rooting 67 systems plays a large role in tree functional response to 68 drought stress Some species develop a 69 deep-rooting system which allows them to maintain high 70 rates of transpiration by using water from deep soil layers 71 as superficial layers dry out (e.g. Zapater et al. 2013) Other 72 species mainly rely on a superficial rooting system, and 73 they suffer sooner from soil water exhaustion as drought 74 conditions worsen. However, a shallow rooting pattern can 75 also sometimes translate into a competitive advantage since 76 the species can absorb water from summer rainy episodes 77 more efficiently 78 These different strategies have been widely studied, but 79 less is known about how the interactions among species 80 with contrasting water-use strategies during drought stress 81 affect their physiological responses to water depletion in 82 natural forest ecosystems. Since the early nineteenth cen- 83 tury, ecologists have been trying to understand how eco- 84 system processes are influenced by community assembly 85 and species interactions In many cases, studies have shown 86 that species interactions can be beneficial for ecosystem 87 functions and services involving the forest carbon and 88 water cycles (Pretzsch et al. 2013a, b; Zhang et al. 2012), 89 even though a negative influence of species interactions 90 can also be found (e.g. Grossiord et al. 2014a). Different 91 mechanisms of species interactions have been suggested to 92 explain these positive effects: Facilitation among species 93 whereby one species will benefit from the presence of 94 another species can take place in mixed-species ecosystems 95 and have an overall positive effect on ecosystem func- 96 tioning Furthermore, complementarity 97 among co-occurring species can lead to a more effective 98 use of available resources and may also be a driver of 99 higher performance in mixed-species ecosystems (Loreau 100 et al. This mechanism occurs when functional traits 101 enable plant species in the community to exploit resources 102 unavailable to others or to use the same resource at a dif- 103, The major physiological adjustment trees 56 undergo to preserve leaf water status We also calculated the daily 359 relative mean sap flux density (F D% ) for each tree as the 360 ratio of F Dmean to F Dmax in order to compare the decline in 361 transpiration among trees along the drought period, 1983. ,
For conifers, current- 367 and previous-year needles were sampled and bulked. The 368 samples were oven-dried at 65 °C for 48 h at INRA Nancy 369 and finely ground (CB2200, Sodemi, St-Ouen l'Aumône, 370 FR) About 1.0 mg of the powdered material from each 371 tree was weighed out Viroflay, FR) 372 and placed into tin capsules (Elemental Microanalysis 373 Limited, Devon, UK) for carbon isotope composition 374 (d 13 C, %) analysis at the Technical Platform of Functional 375 Ecology (OC 081) at the INRA Forest Ecology and Eco- 376 physiology Unit; an EA-GC/IRMS (Delta S, Finnigan 377 MAT, Bremen, Germany; ±0.2 %) was used for the 378 analyses. Isotopic measurements are reported in the delta 379 notation (d, %) according to the Vienna Pee Dee Bel- 380 emnite (VPDB) standard. 381 Data analyses 382 To test the effect of the period of measurements (P1?P5) on 383 REW min , REW mean and REW max , one-way ANOVAs were 384 used followed by Tukey-type post hoc tests to determine 385 differences among periods. To determine differences 386 among species in their response to drought, we used mixed 387 linear models where the fixed effects of species, period and 388 their interaction were tested on F Dmean with ''tree'' as a 389 random factor. Prior to these tests, we employed Bartlett's 390 tests to confirm homogeneity of variance. To determine the 391 influence of the identity of species in the local neigh- 392 bourhood (neighbourhood), the competition intensity (CI), 393 the size of the trees (DBH) and the drought (period) on the 394 decrease in sap flux density for each species, we used 395 mixed linear models where the fixed effects of neigh- 396 bourhood, CI, DBH and period were tested on F D% with 397 ''tree'' as a random factor. The interactions between 398 neighbourhood and period and between CI and period were 399 also included in the model to determine whether the effects 400 of species interactions and competition intensity changed 401 during the drought. Finally, we used mixed linear models 402, 362 Foliar analyses 363 For each selected tree, we used 18-m-long pruning shears 364 to sample 20?30 fully expanded sunlit leaves (Q. faginea) 365 and three to five 30?40-cm-long branches supporting fully 366 mature, p.455, 2013. ,
the 456 identity of species in the immediate neighbourhood 457 (P = 0.025) and of the period of measurement (P \ 0.001) 458 on F D% , but no effect of CI and of DBH (Table 4) The 459 interactions between the level of species interaction and the 460 period as well as between CI and the period were also 461 significant (P \ 0.001, Table 4). F D% of P. sylvestris trees 462 decreased from, p.464 ,
001) on F D% , but no 467 effect of CI and of DBH (Table 4) Furthermore, the 468 interactions between the level of species interaction and the 469 period (P \ 0.001) as well as between CI and the period 470 (P = 0.005) were also significant (Table 4). F D% of Q. 471 faginea trees increased from P1 to P2 and then slightly 472 decreased until P5 (Fig. 4). Trees interacting P, 2009) 530 which access higher water resources during droughts and 531 allow the tree to maintain stomata open over longer periods 532 (Cochard et al. 1996). Regulation of water loss for Quercus 533 species starts only after a very low value of water potential 534 has been crossed (typically -2.0 MPa, p.529, 1993. ,
but our results suggest that such a threshold value was 537 reached late during the summer, probably after P3 Inver- 538 sely, pines, which have a shallower rooting system than 539 oaks, close their stomata early on during periods of limited 540 water availability in order to avoid very negative leaf water 541 potential values and thus potential risks of vessel cavitation 542 However, it must be noted that the two 543 pine species also present important differences in their 544 strategy to deal with drought stress. Indeed, P. sylvestris is 545 less susceptible to xylem embolism than is P. nigra (Choat 546 et al. 2012). Furthermore, P. sylvestris relies mostly on 547 superficial soil layers to extract water, 535 Leaf and soil water potential values were not measured 536 here, 1998. ,
663 Quiroga for their technical assistance We thank the technical Isotope 664 Platform of INRA Nancy for the carbon isotope analyses. A.F. was 665 supported by JAE-PREDOC from CSIC and co-funded by the 666 European Union (Fondo Social Europeo) This work was conducted in 667 the framework of the ARBRE Laboratory of Excellence project 668 (ANR-12-LABXARBRE-01) supported by the French National 669 Research Agency. The research leading to these results was con- 670 ducted within the FunDiv EUROPE project and has received funding 671 from the European Union Seventh Framework Programme, 2007. ,
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