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IZVORNI I ZNANSTVENI ČLANCI – ORIGINAL SCIENTIFIC PAPERS Šumarski list br. 11–12, CXXXIII (2009), 569-576 UDK 630* 243 (001) PHOTOSYNTHETIC RESPONSE OFYOUNG BEECH (Fagus Sylvatica L.) ON RESEARCH PLOTS IN DIFFERENT LIGHT CONDITIONS FOTOSINTETSKI ODZIV MLADIH STABALABUKVE (Fagus sylvatica L.) NA ODABRANIM PLOHAMAU RAZLIČITIM SVJETLOSNIM UVJETIMA 12 Matjaž ČATER , Primož SIMONČIČ ABSTRACT: In view of evident changes in the reaction of European beech (Fagus sylvaticaL.) to environmental changes, five plots with young trees of the same age were established and studied on natural beech sites. Beech trees were equally distributed along the light gradient and were divided according to light conditions. The parameter used for evaluation of light conditions was the indirect site factor (ISF) obtained by the WinScanopy analysis. Three groups of canopy – light conditions were defined: stand conditions (ISF<20), edge (20 describing dependence between intercellular CO2concentration in leaves and assimilation rate (A-Ci) were measured under the same fixed parameters (temperature, flow and CO2concentration, humidity, and light intensity) with Li-6400, to compare responses between different light categories and different plots within comparable light conditions. Differences between canopy, edge and open area responses were confirmed with high significance on all plots as well as between studied forest complexes. On plots from Kočevje region, young beech indicated more shade tolerance, the response to increased light intensity and different CO2concentration was greater than the response of young beech on Pohorje plots within the same light intensities. Responses of trees on plots in managed and virgin forest were also different: young beech response in virgin forest plot was more shade-tolerant, compared to response of young beech from plots in managed forest. Key words: Beech, photosynthesis, light, CO, response 2 INTRODUCTION – Uvod The more frequent and intensive pressures to which cies predominate; in particular natural beech forests forests are exposed are connected with an increasing (Fagus sylvatica L.) (Kutnar 2003). The quality of number of extreme events and consequently higher existing and future beech forests is closely connected risk-rates of forest management decisions, especially on with our understanding of tree-response to different marginaland extreme sites.The importance of autocht-light conditions, especially in an environment of reduhonous tree species in preserving dynamic equilibrium ced light intensity under a mature canopy and in younger and stability in forest ecosystems is frequently emphasi-development stages. Such knowledge leads to correct zed (Zerbe 2002, Hannahet al. 1995, Stanturf and well-tuned spatial and temporal silvicultural measuandMadsen2002). In Slovenia, where forests cover res, which may vary among different silvicultural syover 60 % of the country, sites of mixed broadleaf spe-stems. It is also directed at sustainable development and a better future quality of forests (Kazda 1997). Solar 1 Dr. Matjaž Čater radiation, temperatures and precipitation which inf 2 Dr. Primož Simončič luence the distribution of plants are getting in times of Slovenian Forestry Institute,Večna pot 2, 1000 Ljubljana, matjaz.cater@gozdis.si intensive climatic extremes and climatic changes a new |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 dimension. Evident changes in distribution of plants and species diversity consequently affect primary producti vity (Callaghan etal. 2004). Several key questions about the future response of beech to expected changes such as temperature increase, redistribution of precipitation and increase of atmospheric CO concentration remain open and without answers. Quotations in literature and research results are in most cases unclear and sometimes even contradictory (Po or ter 1998,Lloyd andFarquhar 1996). Photosynthe sis, the first estimate of net productivity could be measured as the response of plants to different light intensity or the response to different concentration of CO 2 which is entering the system in the controlled environment. In spite of good understanding of processes of carbon dynamics at leaf level in a changed CO 2 environment, it is difficult to make a prognosis of the fu ture response of the whole plant, also because of the short time interval of observations and numerous possible interactions that have not yet been recognized.According toBatič(2007) most changes by the increased amount of atmospheric CO could be expected for C3 2 plants at the beginning of saturation curves, especially for the plants that grow in reduced or minimal light conditions, close to compensation point.Wemay therefore expect most changes in shade tolerant species. The research goal was to define range of photosynthetic response in young beech in dependence of light intensity and different concentration of CO between 2 three canopy conditions (shelter, forest edge and gap) on different forest sites. MATERIALAND METHODS – Materijali i metode Research was performed on 10–15 year old beech trees at five selected natural forest stands: at Kladje and Brička in the Pohorje area, atVrhovo and the karstic- Table 1 Research plots characteristics Tablica 1.Značajkepokusnih ploha dinaric area in Kočevski Rog –at Snežna jama (managed forest) and Rajhenav (virgin forest) (Table 1). Plot Ploha Altitude Nadmorska visina (m) Lat (o) Long (o) Annual precipitation Oborina (mm) Annualaverage air T Prosječna godišnja T(oC) Soiltype Tip tla Growing stock Drvna zaliha (m 3 /ha) Brička 1093 46o28’40’’ 15o15’40’ 1190 9,1 Dystric Cambisol 477 Kladje 1308 46o28’48’’ 15o23’24’’ 1066 9,2 Dystric Cambisol 390 Vrhovo 273 45o48’25’’ 15o18’11’’ 1138 9,4 Acric Luvisol 479 Sn. jama 875 45o39’15’’ 15o01’40’’ 1330 8,3 Rendzic Leptosol 612 Rajhenav 865 45o39’36’’ 15o03’36’’ 1330 8,3 Rendzic Leptosol 992 Both Brička and Kladje belong to the acidophilous beech forest type Luzulo albidae-Fagetum (Urbančič and Kutnar 2006) while Snežna jama and Rajhenav belong to dinaric silver fir and beech forest type Omphalodo- Fagetum (Kutnar andUrbančič2008). At each location a research plot was established 100x100m in size, reaching from complete closure to open sky conditions on all plots with little or no exposure. The gradient of natural light conditions was obtained by selecting young trees under a range of canopy openness. On each fenced plot, for 24 young beech trees in comparable light-intensity conditions, their potential light environment was estimated with hemispherical photos (Anonymous 2003). Fine tuning was applied after pilot analysis, so that the light conditions on all plots were comparable.The parameter used for evaluation of light conditions was the indirect site factor (ISF) (Wagner 1994), which is the relative proportion of diffuse light intensity above a defined plant compared to open/gap conditions, (without shading) in percentage (%). Photos were taken with a digital Nikon Coolpix 990 and calibrated fish-eye lens and analyzed withWinScanopy software. In the process of hemispherical photo analysis the vegetation period was defined for each plot group separately; for the diffuse light distribution a “Standard overcast sky” (SOC) model was applied. For the calculation within the vegetation period, the sun’s position was specified every ten (10) minutes.The solar constant was defined as 1370W/m2, 0.6for atmospheric transmissivity and 0.15 for the proportion of diffuse radiation compared to calculated direct potential radiation.According to light conditions three groups were defined: stand conditions (ISF<20), edge (20 of trees on plots ranged from 40 - 70 cm under stand conditions, from 70–110 cm under edge conditions and from 110–220 cm in open area conditions. In each group, four trees were randomly selected for measurement of photosynthesis. In the same leaves nitrogen 2 concentration [mg/cm] was determined to compare macronutrient status in different light categories (Leco CNS-2000 analyzer) (Anonymous,2007). |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 Light saturation curves were established to define comparable ecophysiological response of net assimilation (A) in beech leaves to different light intensities in different plots and in comparable potential light conditions, as described byPotočić etal. (2009).All photosynthesis measurements were performed at a constant temperature of the measurement block (20oC), a CO concentration of 350 µmol/l, flow 500 µmol/s 2 and different light intensities: 0, 50, 250, 600 and 2 1200µmol/ms. Measurements started at ambient light conditions that were reduced to reach zero, then followed by a gradual increase toward maximum values, so that stomata could adapt. A-Ci curves were established to compare and define assimilation response of trees (A) to different intercellular CO concentrations (Ci): measurements were per 2 2 formed at constant light 600 µmol/ms, humidity, constant block temperature 20 oC and flow 500µmol/s, while ambient CO was varied as 0, 50, 100, 350, 700 2 and 1000 µmol/l. Maximal assimilation (A ) rates max and calculated compensation points (CP) for the light saturation andA-Ci curves were used in comparisons of trees between different plots. Both types of response were measured with an LI-6400 portable system on at least three sun leaves per plant, located in the upper third of the tree-crown height on every plot. Twelve trees were measured on each plot, four per same canopy light conditions. Water use efficiency of photosynthesis (WUE), a quantitative measure of the instantaneous gas exchange in leaves was expressed as the ratio of carbon gain per water lost [mol HO/µmol CO] (Larcher 1995, 22 Lambersetal., 1998), while photosynthetic-use efficiency (PNUE) as the carbon gain per unit leaf nitrogen [µmol CO/gN] (Larcher 1995, Lambers et al., 2 1998) for each light category, respectively.Atotal of 20 leaves were sampled per seedling in the upper crown position, then cool-stored in airtight conditions. Fresh leaves were weighed and scanned for the leaf area. Leaves were dried at 105ofor 24 hours until constant weight and weighed for the dry mass. Analyses of variance (ANOVA) and post hoc LSD analysis were used after testing data to meet conditions of normality. Probability values of P<0.05 (*), P<0.01 (**) and P<0.001 (***) were considered significant. Statistical data analysis was done with the programme R(http://www.r-project.org/). RESULTS – Rezultati istraživanja 2 The nitrogen content defined per leaf unit (mg/cm) was different between Pohorje and Kočevje plots in canopy (df ; F=105.13***), edge (df ; F=6.19*) and 1,30 1,30 gap conditions (df ; F=40.99***). On every plot, the 1,30 amount was highest in forest gap and lowest under shelter conditions, except in virgin forest, with maximum values at the forest edge (Table 2). Differences between edge and open area conditions were not significantly different on both plots from Kočevje (Table 3). Table 2 Average leaf nitrogen content per leaf area, water use efficiency (WUE) and photosynthetic nitrogen use efficiency (PNUE) on plots (means ± SE, n=8) Tablica 2.Prosječni sadržaj dušika po jedinici površine lista, efikasnost uporabe vode (WUE) i fotosintetska efikasnost uporabe dušika (PNUE) na plohama (sredine± SE, n=8) Nitrogen (N) [mg/cm 2 ] WUE [mol H 2 O/µmol CO 2 ] PNUE [µmol CO 2 /gN] Plot Ploha Canopy Zastor Edge Rub Gap Otvoreno Canopy Zastor Edge Rub Gap Otvoreno Canopy Zastor Edge Rub Gap Otvoreno Brička 7.5±1.2 9.7±1.3 12.7±1.4 20.0±2.1 18.7±2.3 16.6±2.8 0.04±0.002 0.03±0.002 0.04±0.004 Kladje 8.8±1.0 12.4±1.7 18.2±1.9 22.9±2.4 20.4±1.5 15.8±2.2 0.04±0.005 0.03±0.003 0.02±0.003 Vrhovo 4.5±0.9 8.6±1.5 10.9±2.1 17.9±1.8 12.2±2.7 8.9±1.8 0.06±0.008 0.06±0.003 0.04±0.005 Sn. jama 4.7±0.8 10.4±0.7 10.7±1.6 19.9±3.3 18.8±3.2 15.2±1.9 0.10±0.006 0.08±0.002 0.08±0.003 Rajhenav 3.4±0.8 9.0±0.6 8.9± 1.3 20.3±2.6 20.9±3.1 19.9±3.3 0.08±0.006 0.06±0.002 0.04±0.006 The values for water use-efficiency (WUE) were highest under shelter on all plots, ranging from 17.9 22.9mol HO/µmol CO, with the exception Rajhenav, 22 where maximum values were measured at the forest edge (20.9 mol HO/µmol CO) (Table 2). Although the 22 value measured at the forest edge in Rajhenav was greater from that measured in gap it was statistically not significant (20.9 mol HO/µmol CO compared to 22 20.3mol HO/µmol CO, respectively).Asimilar rela 22 tion was determined for photosynthetic nitrogen use efficiency (PNUE), highest under shelter at Snežna jama (0.10µmol CO/gN).The highest values for the maxi 2 mum assimilation rate (A ) was measured in the open max (gap) atVrhovo, followed by the plots from the Pohorje complex (Brička and Kladje), while the lowest values were measured on plots in Kočevski Rog (Snežna jama, Rajhenav) (Table 4). |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 Table 3 Differences in leaf nitrogen between categories on plots (AVAR and post hoc LSD analysis): 1- shelter; 2 - edge; 3 - open light conditions; NS… non-significant differences Tablica 3.Razlike u sadržaju dušika u lišću izmedju različitih kategorija na plohama (AVAR i post hoc LSD analiza): 1- zastor; 2 - rub; 3 - otvoreno; NS… nesignifikantne razlike df (2, 21) Plot Ploha F p LSD Brička 37.173 0.000 1-2 p=0.0021 1-3 p=0.0000 2-3 p=0.0001 Kladje 93.225 0.000 1-2 p=0.0000 1-3 p=0.0000 2-3 p=0.0000 Vrhovo 70.205 0.000 1-2 p=0.0000 1-3 p=0.0000 2-3 p=0.0001 Sn. Jama 68.914 0.000 1-2 p=0.0000 1-3 p=0.0000 2-3 p=0.7419 NS Rajhenav 88.634 0.000 1-2 p=0.0000 1-3 p=0.0000 2-3 p=0.7419 NS Differences between canopy, edge and open area responses were confirmed with high significance on all plots (Table 5) except in Rajhenav (virgin forest), where no differences between canopy and edge area conditions (df ; F=0.13; NS) were confirmed. 1,14 The response to maximum light in gap conditions between Snežna jama and Rajhenav showed also no differences. Assimilation responses to light were higher in all categories in the virgin forest. The calculated light compensation point (LCP) for edge and gap conditions in Rajhenav and Snežna jama were practically the same 2 (20µmol/ms), values on other plots followed maximal assimilation rates, respectively (data not shown). The assimilation response of young beech between the two forest complexes was also significantly different between canopy (df ; F=285.99***), edge (df ; 1,30 1,30 F=171.68***) and gap conditions (df ; F=93.30***). 1,30 Table 4 Average values of maximum assimilation rates (A ) (means ± SE, n=24) max Tablica 4.Prosječne vrijednosti maksimalne asimilacije (Amax) (sredine ± SE, n=24) Table 5 Differences in maximum assimilation rates (A ) max on plots (AVAR and post hoc LSD analysis): 1- shelter; 2 - edge; 3 - open light conditions; NS… non significant differences Tablica 5.Razlike u maksimalnoj asimilaciji (Amax) na plohama (AVAR i post hoc LSD analiza): 1- zastor; 2 - rub; 3 - otvoreno; NS… nesignifikantne razlike df (2, 21) Plot Ploha F p LSD Brička 58.681 0.000 1-2p=0.0000 1-3 p=0.0000 2-3 p=0.0001 Kladje 28.804 0.000 1-2 p=0.0002 1-3 p=0.0000 2-3 p=0.0049 Vrhovo 442.675 0.000 1-2 p=0.0000 1-3p=0.0000 2-3 p=0.0000 Sn. Jama 75.266 0.000 1-2 p=0.0000 1-3 p=0.0000 2-3 p=0.0000 Rajhenav 12.495 0.000 1-2 p=0.6002 NS 1-3p=0.0002 2-3 p=0.0006 Maximum assimilation values forA-Ci curves (A max ) (Table 6) showed similar reaction of trees as in the case of maximum assimilation values measured for the light curves (A ) (Table 4). A-Ci max The response of young beech was greatest in the category of canopy gap and lowest under shelter on all plots.The highest response to increased CO concentra 2 tion was evident atVrhovo, followed by plots on Pohorje (Brička and Kladje) and lowest in Kočevski Rog (Rajhenav, Snežna jama). Comparison of the calculated compensation point for CO between light catego 2 ries showed no significant differences (data not shown). Differences between forest complexes were statistically different in all categories: canopy (df ; 1,30 F=6.47**), edge (df ; F=13.17**) and gap conditions 1,30 (df ; F=33.41***). 1,30 Table 6 Maximum assimilation values for A-Ci curves (A ); (means ± SE, n=24 trees). max A-Ci Tablica 6.Maksimalna asimilacija (Amax A-Ci); (sredine ± SE, n=24 stabla). Amax (µmol /m 2 s) Canopy Zastor Edge Rub Gap Otvoreno Brička 7.3±0.4 9.8±0.8 11.9±1.1 Kladje 8.3±0.3 9.7±0.5 10.7±0.9 Vrhovo 6.1±0.4 9.3±0.4 13.2±0.6 Snežna jama 4.8±0.4 6.5±0.5 8.0±0.7 Rajhenav 7.1±0.3 7.2±0.5 8.2±0.6 Amax A-Ci (µmol /m 2 s) Canopy Zastor Edge Rub Gap Otvoreno Brička 6.1±0.7 7.7±0.6 12.3±0.5 Kladje 8.8±0.6 9.6±0.5 10.4±0.6 Vrhovo 5.4±0.4 7.1±0.8 13.4±1.0 Snežna jama 3.4±0.4 5.4±0.4 9.4±0.6 Rajhenav 7.8±0.4 8.3±0.6 9.7±0.6 |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 In the Kočevski Rog area, differences among the tree ged one (at Snezna jama).Assimilation rates were higlight categories are smaller in the virgin forest where no hest in all categories in virgin forests, despite the comdifferences in response between canopy and edge were parable amount of nitrogen in leaves on the two plots. confirmed (Post hoc LSD, p=0.0969) than in the mana- DISCUSSIONAND CONCLUSIONS – Rasprava i zaključci Sensitivity of photosynthesis is similar for all C 3 plants and is in proportion with mesophyll CO concen 2 tration (Farquhar etal. 1980). Many studies indicate that trees have higher mesophyll resistance for CO, 2 consequently lower photosynthesis and are therefore more susceptible to the increase of atmospheric CO 2 concentration. In the view of climatic changes numerous and often contradictory conclusions are being presented about the response of plants and future development of tree adaptation to environmental changes, especially due to temperature increase (Körner, 2006), decrease in amount of precipitation and changes in water supply (Davies 2006) and increase of atmospheric CO con 2 centration (Ziska andBunce2006). Light, nutrients, water and CO are abiotic parame 2 ters, necessary for the plant growth. Efficiency and photosynthetic regulation are governed by ribulose-1,5 biphosphat carboxylase (rubisco), which is genetically defined (Cheng etal. 1998). In general, by higher atmospheric CO; protein synthesis in leaves increases, 2 stomatal aperture decreases, water use-efficiency and C/N relation on the leaf level are increased, while on the whole plant level growth is stimulated (Kimball 1993, Ghannoum et al. 2000). By the increased amount of CO photosynthesis per 2 unit of the leaf area would increase, which is dependent on the nitrogen supply. Respiration and root activities would also increase, while biomass would be allocated into roots (sweet chestnut) or increased proportionally over whole plant (beech), which indicates a species- specific response (Kohen etal. 1993). In spite of relatively good insight into processes of carbon dynamics on the leaf level in changed CO en 2 vironment it is difficult to make a prognosis of future response of the whole plant also because of a short- time interval of observations and numerous possible interactions that haven’t been recognized yet (Increased WUE might stimulate development of foliar fungi (Thompson andDrake1994) while more sugars in assimilation apparatus might stimulate the development of pathogens and infections (Hibberd et al. 1996) etc.). Recent research quote up to 30 % increase of growth in ambient with two times higher CO envi 2 ronment (Medylin etal. 2001).Asmaller probability that such increase would reflect in long term growth in assimilation was confirmed by Batič 2007, where growth only increased at the beginning, and was later reduced in time. Our analysis conirmed the differences in response between beech under shelter, at the forest edge and in the open. In spite of these differences, the highest assimilation rates were measured on the research plots of Pohorje complex and lowest on the plots of Kočevski Rog. Results also indicate a different response of young beech between managed forest (Snežna jama) and virgin forest (Rajhenav); differences between light categories were more pronounced in the managed forest, while in virgin forest response to same light conditions was more intense than in managed forest. Photosynthetic yield in all categories was higher in virgin forest. Light compensation point was higher on plots of Pohorje complex compared to plots in Kočevski Rog (data not shown). Water use efficiency (WUE) was in all cases highest under shelter and lowest in open conditions, similar to photosynthetic nitrogen use-efficiency (PNUE). Self shading and nitrogen redistribution within whole plant could potentially underlie the degree of photosynthetic acclimation to elevated CO (Takeu 2 chi et al. 2001); it is clear that interactions with other potential environmental variables (light, nutrients) will determine the regulation of carbon sources and sinks at the leaf level (Lewis etal. 2002), but the ability to utilize the knowledge and predict a whole plant response is still limited and subject of controversy (Poorter 1998, LloydandFarquhar1996). There were no significant differences in the content of leaf nitrogen between plots. Leaf nitrogen values were on all plots (expressed in units per leaf area) highest in open conditions, without shading. Response of young beech to different CO concentrations was simi 2 lar to response of young beech to different light intensity; differences between managed and virgin forest were even bigger under canopy and edge conditions. In Kočevski Rog young beech is more shade tolerant, relative response to increased light intensity and different CO concentration is higher than response of young 2 beech in Pohorje within same light intensities. Responses in managed and virgin forest are different: in the virgin forest young beech trees are more shade-tolerant, reaction of different light categories to elevated CO concentration is similar and more homogenous, compared to managed forest where differences between cate gories are more pronounced. Kočevje region (Snežna jama and Rajhenav) is well known for its forest manage |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 ment, with a long tradition of a sustainable and close-tonature approach, with a single-tree selection method (Diaci 2006). In contrast, in the Pohorje complex, with potential beech sites, Norway spruce has been favored in the last century and nowadays beech is gradually replacing spruce either by underplanting or by natural regeneration (Diaci 2006). In case of the response in young beech trees, differences in assimilation rate may reflect not only the different forest management history, but also a different genetic background. ACKNOWLEDGEMENT – Zahvala Autors wish to thank dr. Nenad Potočić, from Šumarski Institut Jastrebarsko, Croatia, forconstructive suggestions on the manuscript text and revision of titles and abstract in Croatian language.The article is a result of the “L4-6232; Carbon dynamic in natural beech forest” project, financed by the Slovenian Research Agency and Programme research group. REFERENCES – Literatura Anonymous,2007. CE CLRTAPICPonAssessment and Monitoring ofAir Pollution Effects on Forests MANUAL on methods and criteria for harmonized sampling, assessment, monitoring and analysis of the effects of air pollution on forests, Part IV, Sampling andAnalysis of Needles and Leaves, Updated 05/2005 (page 7: 2007); (http://www.icp-forests.org/pdf/manual4.pdf) Anonymous, 2003. WinSCANOPY2003b for hemispherical image analysis, Regent instruments inc., Manual, 106 p. Batič, F., 2007.Rastline in podnebne spremembe = Plants and climate change. In: Jurc, Maja (ed.). Podnebne spremembe : vpliv na gozd in gozdarstvo : impact on forest and forestry, (Studia forestalia Slovenica, št. 130). Ljubljana: Biotehniška fakulteta, Oddelek za gozdarstvo in obnovljive gozdne vire: = Biotechnical Faculty, Department of Forestry and Renewable Forest Resources Slovenia, 2007, p. 51–66. Callaghan, T.V., L.O. Bjorn, Y. Chernov, T. Chapin, T.R. Christensen, B. Huntley, R.A.Ims, M.Johansson, D.Jolly, S.Jonasson, N. Matveyeva, N. Panikov, W. Oechel, G. Shaver, J. Elster, I.S. Jonsdottir, K. Laine, K. Taulavuori, E. Taulavuori & C. Zocker, 2004. Responses to projected changes in climate and UV-B at the species level,Ambio, 33, (7), p.418–435. Davies, W.J., 2006. Responses of plant growth and functioning to changes in water supply in a changing climate. In: Plant Growth and Climate Change, J.I.L. Morison (Ed.), M.D. Morecroft (Ed.),Wiley-Blackwell, pp 96–117. Diaci,J., 2006. Fifty years of restoration in norway spruce replacement forests in Slovenia. In: Simončič, P. (ed.), Čater, M. (ed.). Splošne ekološke in gozdnogojitvene osnove za podsadnjo bukve (Fagus sylvatica L.) v antropogenih smrekovih sestojih, (Studia forestalia Slovenica, št. 129). Ljubljana: Gozdarski inštitut Slovenije, Silva Slovenica, 2006, p. 130–141. Cheng,S.-H., B.D.Moore,J.R.Seemann,1998. Effects of short- and long-term elevated CO on 2 the expression of Ribulose-1,5 biphosphate carboxylase/ oxygenase genes and carbohydrate accumulation in leaves ofArabidopsis thaliana (L.) Heynh. Plant Physiology, 116, p.715–723. Farquhar,G.D., S. von Caemmerer, and J.A. Berry, 1980. A Biochemical Model of Photosynthetic CO2 Assimilation in Leaves of C3 species. Planta149, p. 78–90. Ghannoum, O., S. von Caemmerer, L.H. Zis ka, J.P. Conroy,2000. The growth response of C plants to rising atmospheric CO partial 4 2 pressure: a reassessment. Plant Cell Environm., 23, p. 931–942. Hannah,L., J.L. Carr,A. Lankerani, 1995. Human disturbance and natural habitat: biome level analysis of a global data set. Biodiv. Conserv. 4, p. 128–155. Hibberd,J.M., R.Whitbread, J.F.Farrar,1996. Effect of elevated concentrations of CO on in 2 fection ofbarley byErysiphe graminis. Physiol. Mol. Plant. Pathol., 48, p. 37–49. Kazda,M., 1997. Lichtverteilung inWaldbeständen - Konsequenzen für denWaldbau. Österreichische Forstzeitung, p. 11–13. Kimball,B.A., 1993. Effects of increasing atmospheric CO on vegetation.Vegetatio, 104/105, p. 2 65–83. Kohen,A.E.L., L. Venet, M. Mousseau, 1993. Growth and photosynthesis of two deciduous forest species at elevated carbon dioxide, Functio nal Ecology, 7, p. 480–486. Kutnar,L., 2003. Forest vegetation of Slovenia. In: Intensive monitoring programme in Slovenia |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 čimbenik (ISF), dobiven analizom hemisfernih snimaka pomoću sustava Win- Scanopy: zastor krošanja (ISF<20), rub sastojine (20 fotosintetskog kapaciteta, krivulje svjetlosnog zasićenja (0, 50, 250, 600 i 1200 µmol/m2s) i A-Ci krivulje (0, 100, 400, 700 i 1000 µmol CO2/l) dobivene su pomoću Li-Cor LI-6400 u kontroliranom okruženju (temperatura, protok i koncentracija CO2, zračna vlaga). Analize sadržaja dušika u lišću napravljene su Leco CNS-2000 analizatorom. Potvrđene su signifikantne razlike u reakciji mladih bukava između odabranih kategorija, kao i između različitih šumskih kompleksa. Mlade bukve na plohama iz Kočevskog pokazale su veću toleranciju na sjenu, a odziv na porast koncentracije CO2je pri istim intenzitetima osvijetljenosti bio veći nego kod mladih bukava iz Pohorskog kompleksa. Odziv mladih bukava bio je sig ni fi kant no različit između prašume (Rajhenav) i gospodarske šume unutar istog šumskog kompleksa: odziv u prašumi pokazuje veću toleranciju na sjenu. Ključne riječi:Bukva, fotosinteza, svjetlo, CO2, odziv |
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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576 (IMP-SI): basic structural document : project do cument. Ljubljana: Slovenian Forestry Institute; Wageningen: Alterra, 2003. p. 21 Kutnar,L., M.Urbančič,2008. Influence of site and stand conditions on diversity of soil and vegetation in selected beech and fir-beech forests in the Kočevje region. Zbornik gozdarstva in lesarstva 80, p. 3–30. Körner, C., 2006. Significance of temperature in plant life. In: Plant Growth and Climate Change, J.I.L. Morison (Ed.), M.D. Morecroft (Ed.), Wiley-Blackwell, pp 48–69. Lambers,H., F.S. Chapin,III, T.L. Pons,1998. Plant Physiological Ecology.- Springer, New York – Berlin – Heidelberg – Barcelona – Budapest – Hong Kong – London – Milan – Paris – Singapore –Tokyo, 550 pp. Larcher, W., 1995. Physiological plant ecology.Ecophysiology and stress physiology of functional groups.-Berlin, Springer-Verlag, 506 p. Lewis,J.D., X.Z. Wang,K.L.Griffin, D.T.Tissue, 2002. Eeffects of age and ontogeny on photosynthetic responses of a determinate annual plant to elevated CO cocentrations. Plant 2 Cell Environm., 25, p. 359–368. Lloyd,J., G.D. Farquhar,1996. The CO depen 2 dence of photosynthesis, plant growth responses to elevated atmospheric CO concentrations and 2 their interaction with soil nutrient status. I. Ge neral Principles and Forest Ecosystems., Funct. Ecol., 10, p.4–32. Medlyn,B.E., A.Rey, C.V.M.Barton & M.Forstreuter, 2001.Above-ground growth responses of forest trees to elevated atmospheric CO 2 concentrations. In:The Impact od Carbon Dio xide and Other Greenhouse Gases on Forest Ecosystems (eds D.F. Karnosky, R. Ceulemans, G.E. Scarascia-Mugnozza & J.L. Innes) CABI Publishing, NewYork, pp. 127–146. Poorter,H., 1998. Do slow-growing species and nutrient- stressed plants respond relatively strongly to elevated CO? Global Change Biol., 4., p. 2 693–697. Potočić,N., I.Seletković, M.Čater,T.Ćosić, M. Šango, M. Vedriš, 2009. Ekofiziološki odziv suncu izloženih sadnica obične bukve (Fa gus sylvaticaL.) pri različitim razinama gnojidbe: Ecophysiological Response of Sun-Exposed Com mon Beech (Fagus Sylvatica L.) Seedlings under Different Fertilization Levels. Šumarski list br. 5–6, p. 289–300. Stanturf,J.,A., P.Madsen,2002. Restoration concepts for temperate and boreal forests of North America andWestern Europe. Plant Biosystems 136, p. 143–158 Takeuchi, Y., M.E. Kubiske, J.G. Isebrands, K.S.Pregtizer, G.Hendrey, D.F.Karnosky, 2001. Photosynthesis, light and nitrogen relationships in a young deciduous forest canopy under open-air CO enrichment. Plant cell Envi 2 ron., 24, p. 1257–1268. Thompson, G.B., B.G. Drake, 1994. Insects and fungi on a C sedge and a C grass exposed to 34 ele vated atmospheric CO concentrations in 2 open-top chambers in the field. Plant Cell Environm., 17, p. 1161–1172. Urbančič,M., L.Kutnar,2006. Site conditions of the plot Brička and comparisons with other Sustman plots. In: Simončič, P. (ed.), Čater, M. (ed.). Splošne ekološke in gozdnogojitvene osnove za podsadnjo bukve (Fagus sylvatica L.) v antropogenih smrekovih sestojih, Studia forestalia Slovenica, 129, Ljubljana, Slovenian Forestry Institute, p.68–85. Zerbe,S., 2002. Restoration of natural broad-leaved woodland in Central Europe on sites with coniferous forest plantations. For. Ecol. Manage. 167, p. 27–42. Wagner,S., 1994. Sthrahlungschätzung inWäldern durch hemisphärische Fotos – Methode undAnwendung, Berichte des Forschungszentrums Waldökosysteme, Reihe A, Bd 123, Göttingen, 166 p. Ziska,L.H., J.A. Bunce,2006. Plant responses to rising atmospheric carbon dioxide. In: Plant Growth and Climate Change, J.I.L. Morison (Ed.), M.D. Morecroft (Ed.), Wiley-Blackwell, pp 17–47. SAŽETAK: Glede uočenih promjena u reakciji bukve (Fagus sylvaticaL.) u odnosu na ekološke promjene, odabrano je pet ploha mlade bukve jednake starosti na prirodnim staništima, koje su bile jednakomjerno raspoređene na svjetlosnom gradijentu od zastora odrasle sastojine, šumskog ruba do svjetlosnih uvjeta na otvorenome. Kriterij za grupiranje bio je neizravni stanišni |