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1.
We examined the principal differences in photosynthetic characteristics between sun and shade foliage and determined the relative importance of biochemical and stomatal limitations during photosynthetic induction. Temperate-zone broadleaf and conifer tree species, ranging widely in shade tolerance, were investigated from one locality in the Czech Republic. The study species included strongly shade-tolerant Abies alba Mill. and Tilia cordata Mill., less shade-tolerant Fagus sylvatica L. and Acer pseudoplatanus L. and sun-demanding Picea abies (L.) Karst. In the fully activated photosynthetic state, sun foliage of all species had significantly higher maximum CO(2) assimilation rates, maximum stomatal conductance and maximum rates of carboxylation than shade foliage. Compared with shade leaves, sun leaves had significantly higher nocturnal stomatal conductances. In all species, shade foliage tended to have higher induction states 60 s after leaf illumination than sun foliage. Sun and shade foliage did not differ in the rate of disappearance of the transient biochemical limitation during the induction phase. Longer time periods were required to reach 90% photosynthetic induction and 90% stomatal induction in sun foliage than in shade foliage of the less shade-tolerant F. sylvatica and A. pseudoplatanus and in sun-demanding P. abies; however, in sun foliage of the strongly shade-tolerant species T. cordata and A. alba, the time needed for photosynthetic induction was similar to, or less than, that for shade foliage. Shade but not sun needles of P. abies and A. alba had significantly slower induction kinetics than the broadleaf tree species. Among species, the sun-demanding P. abies exhibited the shortest stomatal induction times in both sun and shade leaves. Independently of shade tolerance ranking, the transient stomatal and total limitations that characterize photosynthetic induction were relieved significantly earlier in shade foliage than in sun foliage. Sun foliage generally exhibited a hyperbolic photosynthetic induction response, whereas a sigmoidal induction response was more frequent in shade foliage. The different relative proportions of transient biochemical and stomatal limitations during photosynthetic induction in sun and shade foliage indicate an essential role of stomata in photosynthetic limitation during induction, mainly in shade foliage, with a consequent influence on the shape of the photosynthetic induction curve.  相似文献   

2.
Robakowski P 《Tree physiology》2005,25(9):1151-1160
Susceptibility to photoinhibition of the evergreen conifers Abies alba Mill., Picea abies (L.) Karst. and Pinus mugo Turra was investigated in an unheated greenhouse during winter and spring 2003. Photosynthetic performance of the seedlings was assessed by chlorophyll a fluorescence and analyses of chlorophyll and total carotenoid concentrations in needles. During winter months, maximum quantum yield of PSII photochemistry (ratio of variable to maximum fluorescence, Fv/Fm) was significantly greater in A. alba than in P. abies and P. mugo. Abies alba also sustained higher maximum apparent electron transport rate (ETRmax) than P. abies and P. mugo. Total concentrations of chlorophyll and carotenoids in needles decreased during the winter in P. mugo and P. abies, but remained stable in A. alba. For all species, Fv/Fm decreased from December until February and then increased to a maximum in April. Photoinhibition was greatest (Fv/Fm < 0.80) in all seedlings in February, the month with the lowest mean temperature. Saturating photosynthetic photon flux (PPFsat) and ETRmax were positively related to air temperature. All species had lower values of ETRmax and PPFsat in winter than in spring. Non-photochemical quenching of chlorophyll fluorescence (NPQ) was highest at low air temperatures. Differences among species in susceptibility to winter photoinhibition resulted from their specific light preferences and led to different mechanisms to cope with photoinhibitory stress. The more shade-tolerant A. alba sustained a higher photosynthetic capacity in winter than P. abies and P. mugo. Winter photoinhibition in P. abies, P. mugo and, to a lesser extent, in A. alba may reflect adaptive photoprotection of the photosynthetic apparatus in winter.  相似文献   

3.
We investigated susceptibility to photoinhibition in leaves acclimated to different light regimes in intermediately shade-tolerant Japanese oak (Quercus mongolica Fisch. ex Turcz. var. crispula (Blume) Ohashi) and shade-tolerant Japanese maple (Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara), to elucidate adaptability to gap formation in leaves differing in shade acclimation. We hypothesized that there is a tradeoff between shade adaptation and capacity to mitigate photoinhibition associated with leaf morphology. We simultaneously measured chlorophyll fluorescence and gas exchange in seedlings that had been grown in full sunlight (open), 10% of full sun (moderate shade) and 5% of full sun (deep shade). Shade-tolerant A. mono adapted to deep shade through changes in leaf morphology, lowering its leaf mass per area (LMA), but Q. mongolica showed little change in LMA between moderate and deep shade. Photochemical quenching (qP) did not differ between species in full sunlight and moderate shade; however, in deep shade, qP of Q. mongolica was higher than that of A. mono, suggesting that Q. mongolica grown in deep shade is less susceptible to photoinhibition at gap formation. This is consistent with the finding that chronic photoinhibition 3 days after the transfer to full sunlight, indicated by the decrease in maximum photochemical efficiency, Fv/Fm, at predawn, was less in deep-shade-grown Q. mongolica than in deep shade-grown A. mono. In deep shade, the electron transport rate (ETR) of Q. mongolica was higher than that of A. mono, whereas thermal energy dissipation through photosystem II antennae, indicated by non-photochemical quenching, was lower in Q. mongolica than in A. mono. In deep shade, the greater ETR capacity in Q. mongolica in association with higher LMA and higher leaf N content could contribute to maintaining high qP and mitigating photoinhibition. These results indicate that, by maintaining a high electron transport capacity even in deep shade, the gap-dependent and intermediate-shade-tolerant Q. mongolica trades improved shade adaptation for higher growth potential when a gap event occurs.  相似文献   

4.
We investigated the differential roles of physiological and morphological features on seedling survivorship along an experimental irradiance gradient in four dominant species of cool temperate-Mediterranean forests (Quercus robur L., Quercus pyrenaica Willd., Pinus sylvestris L. and Pinus pinaster Ait.). The lowest photochemical efficiency (F(v)/F(m) in dark-adapted leaves) was reached in deep shade (1% of full sunlight) in all species except Q. robur, which had the lowest photochemical efficiency in both deep shade and 100% of full sunlight. Species differed significantly in their survival in 1% of full sunlight but exhibited similar survivorship in 6, 20 and 100% of full sunlight. Shade-tolerant oaks had lower leaf area ratios, shoot to root ratios, foliage allocation ratios and higher rates of allocation to structural biomass (stem plus thick roots) than shade-intolerant pines. Overall phenotypic plasticity for each species, estimated as the difference between the minimum and the maximum mean values of the ecophysiological variables studied at the various irradiances divided by the maximum mean value of those variables, was inversely correlated with shade tolerance. Observed morphology, allocation and plasticity conformed to a conservative resource-use strategy, although observed differences in specific leaf area, which was higher in shade-tolerant species, supported a carbon gain maximization strategy. Lack of a congruent suite of traits underlying shade tolerance in the studied species provides evidence of adaptation to multiple selective forces. Although the study was based on only four species, the importance of ecophysiological variables as determinants of interspecific differences in survival in limiting light was demonstrated.  相似文献   

5.
We hypothesized that photoinhibition of shade-developed leaves of deciduous hardwood saplings would limit their ability to acclimate photosynthetically to increased irradiance, and we predicted that shade-tolerant sugar maple (Acer saccharum Marsh.) would be more susceptible to photoinhibition than intermediately shade-tolerant red oak (Quercus rubra L.). After four weeks in a canopy gap, photosynthetic rates of shade-developed leaves of both species had increased in response to the increase in irradiance, although final acclimation was more complete in red oak. However, photoinhibition occurred in both species, as indicated by short-term reductions in maximum rates of net photosynthesis and the quantum yield of oxygen evolution, and longer-term reductions in the efficiency of excitation energy capture by open photosystem II (PSII) reaction centers (dark-adapted F(v)/F(m)) and the quantum yield of PSII in the light (phi(PSII)). The magnitude and duration of this decrease were greater in sugar maple than in red oak, suggesting greater susceptibility to photoinhibition in sugar maple. Photoinhibition may have resulted from photodamage, but it may also have involved sustained rates of photoprotective energy dissipation (especially in red oak). Photosynthetic acclimation also appeared to be linked to an ability to increase leaf nitrogen content. Limited photosynthetic acclimation in shade-developed sugar maple leaves may reflect a trade-off between shade-tolerance and rapid acclimation to a canopy gap.  相似文献   

6.
In a 4-year study, we investigated changes in leaf physiology, crown morphology and whole-tree biomass allocation in seedlings and saplings of shade-tolerant sugar maple (Acer saccharum Marsh.) and intermediate shade-tolerant yellow birch (Betula alleghaniensis Britt.) growing in natural understory light (0.5 to 35% of full sunlight) or in understory light reduced by 50% with shade nets to simulate the effect of gap closure. Leaf physiological parameters were mainly influenced by the light gradient, whereas crown morphological and whole-tree allocational parameters were mainly influenced by tree size. No single physiological, morphological or allocational trait was identified that could explain the difference in shade tolerance between the species. Yellow birch had higher growth rates, biomass allocation to branches and leaf physiological plasticity and lower crown morphological plasticity in unmodified understory light than sugar maple. Sugar maple did not display significant physiological plasticity, but showed variation with tree size in both crown morphology and whole-tree biomass allocation. When sugar maple was small, a greater proportion of whole-tree biomass was allocated to roots. However, physiological differences between the species decreased with decreasing light and most morphological and allocational differences tended to disappear with increasing tree size, suggesting that many species differences in shade-tolerance are expressed mainly during the seedling stage. Understory trees of both species survived for 4 years under shade nets, possibly because of higher plasticity when small and the use of stored reserves when taller.  相似文献   

7.
5种箬竹属竹种叶绿素荧光特性的比较   总被引:2,自引:0,他引:2  
利用叶绿素荧光成像技术研究了5种箬竹属(Indocalamus Nakai)竹种叶片光合作用的横向异质性,结果表明,吸光系数Abs、PSⅡ最大量子产量Fv/Fm、光化学淬灭qP的异质性较低,非光化学淬灭NPQ的异质性较高。比较稳态荧光和荧光日变化,胜利箬竹的Fv/Fm、PSⅡ实际量子产量Yield、相对电子传递速率ETR都保持相对较高水平,矮箬竹和小叶箬竹则较低。  相似文献   

8.
Kitao M  Lei TT  Koike T 《Tree physiology》1998,18(2):135-140
We used chlorophyll fluorescence to examine photosynthetic responses to excess Mn accumulation in leaves of four tree species differing in successional traits. Betula ermanii Cham. (Be) and Alnus hirsuta Turcz. (Ah) were studied as representatives of early-successional species. Ulmus davidiana Planch. var. japonica (Rehder) Nakai (Ud) was selected as a mid-successional species, and Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara (Am) was chosen as a late-successional species. In Be, Ah and Am, high foliar concentrations of Mn had little effect on maximum photochemical efficiency of photosystem II (PSII), as indicated by the values of dark-adapted F(v)/F(m), whereas a significant decrease was observed in Ud. Photochemical quenching (qP) and the excitation capture efficiency of open PSII (F'(v)/F'(m)) decreased with increasing leaf Mn concentration at photosynthetic steady state after a 15-min exposure to 430 &mgr;mol m(-2) s(-1) PPFD. Compared with early-successional species, these decreases were greater in mid- and late-successional species leading to lower effective quantum efficiencies of PSII (DeltaF/F'(m) = qP x F'(v)/F'(m) = (F'(m) - F)/F'(m)). To determine the extent of photoinhibition, F(v)/F(m) of the illuminated leaves was remeasured after a 15-min dark period. Compared with the dark-adapted F(v)/F(m), we observed a significant decrease in F(v)/F(m) in Am leaves containing high concentrations of Mn. These chlorophyll fluorescence studies indicate that the early-successional species Be and Ah have a higher tolerance to excessive accumulations of Mn in leaves than the mid- and late-successional species Ud and Am.  相似文献   

9.
Mori A  Takeda H 《Tree physiology》2004,24(6):661-670
Light-related plasticity of crown morphology and within-crown characteristics were investigated in understory sun and shade saplings of three codominant subalpine conifers, Abies mariesii M.T. Mast., Abies veitchii Lindl. and Picea jezoensis var. hondoensis (Mayr) Rehd. Compared with those of sun saplings, current-year shoots of shade saplings allocated less biomass to needles, resulting in less dense needle packing and hence less mutual needle shading. The proportion of lateral branch biomass in foliage was either similar in sun and shade saplings or greater in shade saplings, depending on the species, suggesting that, over the lifetime of a branch, greater needle longevity in shade compensates for reduced biomass investment in needles of current-year shoots of shade saplings. Saplings with slower-growing branches tended to have greater needle life spans, suggesting that plasticity of branch growth rate and plasticity of needle life span are interdependent. Both Abies species showed greater light-related plasticity of needle life span and branch growth than P. jezoensis. The greater shade tolerance of the Abies species derives from their broad flattened crowns with slow-growing branches. This type of crown development incurs substantial support costs, but the long needle life span of shade saplings of the Abies species compensates, at least in part, for their low annual investment in foliage, especially in the case of A. mariesii, which has a longer needle life span and slower-growing and stouter branches than A. veitchii. Compared with the Abies species, P. jezoensis had a less plastic crown morphology, and less variability of needle life span and branch growth in response to light, resulting in lower shade tolerance. However, compared with the flattened crown of Abies shade saplings, the conical crown of P. jezoensis saplings imposes a smaller support cost, making this species better adapted to rapid height growth than to survival in shade.  相似文献   

10.
We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.  相似文献   

11.
We assessed the irradiance-related plasticity of hydraulic architecture in saplings of Betula pendula Roth., a pioneer species; Acer pseudoplatanus L., Fraxinus excelsior L. and Quercus robur L., which are post-pioneer light-requiring species; and Quercus petraea Matt. Liebl. and Fagus sylvatica L. Plants were grown in pots in 36%, 16% and 4% of full sunlight. Hydraulic conductance was measured with a high-pressure flow-meter in entire, in situ root systems and in excised shoots. Leaf-specific whole-plant conductance (LSC) increased with irradiance, due, in part, to an effect of irradiance on plant size. In addition, there was a size-independent effect of irradiance on LSC due, in part, to an increase in root hydraulic conductance paralleled by an increase in root biomass scaled to leaf area. Changes in shoot conductivity also contributed to the size-independent plasticity of LSC. Vulnerability to cavitation measured in current-year twigs was much larger in shade-grown plants. Betula pendula had the highest whole-plant, root and shoot conductances and also the greatest vulnerability to cavitation. The other species were similar in LSC, but showed some variation in root conductance scaled to biomass, with Q. robur, Q. petraea and F. sylvatica having the lowest root conductance and susceptibility to cavitation. All species showed a similar irradiance-related plasticity in LSC.  相似文献   

12.
Grassi G  Bagnaresi U 《Tree physiology》2001,21(12-13):959-967
The role of morphological versus physiological foliar plasticity in the capacity for, and mechanisms of, photosynthetic acclimation was assessed in Picea abies (L.) Karst. and Abies alba Mill. saplings in a forest gap-understory light gradient (relative irradiance, RI, ranging from 0.02 to 0.32). The species investigated showed a similar foliar morphological plasticity along the light gradient, at both the needle level (through alteration in leaf dry mass per area) and the shoot level (through alteration in the silhouette area ratio, e.g., shoot silhouette to projected needle area ratio). In both species chlorophyll (Chl) concentration on a mass basis decreased at increasing RI, but was independent of RI when expressed on an area basis. In contrast, leaf N concentration on a mass basis was independent of RI, but was positively influenced by RI when expressed on an area basis. The parameters describing photosynthetic performance at low light (dark respiration rate, apparent quantum yield and light compensation point) suggest that Abies alba was better suited to maintain a positive carbon balance in shaded conditions. By contrast, parameters describing biochemical capacity at high light (maximum electron transport rate, Jmax and maximum ribulose-1,5-biphosphate carboxylation capacity, Vcmax) indicate that only Picea abies was capable of acclimating physiologically to high photosynthetic photon flux densities (PPFDs) by increasing nitrogen partitioning to Rubisco and Vcmax/mass by increasing RI. These results support the hypothesis that interspecific differences in nitrogen partitioning within the photosynthetic apparatus may provide a mechanistic basis for species separation along a light gradient. The differences in photosynthetic plasticity observed are likely to influence regeneration patterns and habitat breadth of the species investigated. The limited ability of Abies alba saplings to exploit high-light conditions may be a competitive disadvantage in large canopy gaps and thus limit recruitment of this species to small gaps.  相似文献   

13.
Prunus domestica L. has green leaves, whereas Prunus cerasifera Ehrh. var. atropurpurea has red leaves due to the presence of mesophyll anthocyanins. We compared morphological and photosynthetic characteristics of leaves of these species, which were sampled from shoots grafted in pairs on P. domestica rootstocks, each pair comprising one shoot of each species. Two hypotheses were tested: (1) anthocyanins protect red leaves against photoinhibition; and (2) red leaves display shade characteristics because of light attenuation by anthocyanins. Parameters were measured seasonally, during a period of increasing water stress, which caused a similar drop in shoot water potential in each species. As judged by predawn measurements of maximum PSII yield, chronic photoinhibition did not develop in either species and, despite the anthocyanic screen, the red leaves of P. cerasifera displayed lower light-adapted PSII yields and higher non-photochemical quenching than the green leaves of P. domestica. Thus, it appears that, in this system, anthocyanins afford little photoprotection. As predicted by the shade acclimation hypothesis, red leaves were thinner and had a lower stomatal frequency, area- based CO2 assimilation rate, apparent carboxylation efficiency and chlorophyll a:b ratio than green leaves. However, red leaves were similar to green leaves in conductivity to water vapor diffusion, dry-mass-based chlorophyll concentrations and carotenoid:chlorophyll ratios. The data for red leaves indicate adaptations to a green-depleted, red-enriched shade, rather than a neutral or canopy-like shade. Thus, green light attenuation by anthocyanins may impose a limitation on leaf thickness. Moreover, the selective depletion of light at wavelengths that are preferentially absorbed by PSII and chlorophyll b may lead to adjustments in chlorophyll pigment ratios to compensate for the uneven spectral distribution of internal light. The apparent photosynthetic cost associated with lost photons and reduced leaf thickness, and the absence of a photoprotective advantage, suggest that there are other, yet to be identified, benefits for permanently anthocyanic leaves of P. cerasifera.  相似文献   

14.
Photoinhibition of photosynthesis and photosynthetic recovery were studied in detached needles of cypress (Cupressus sempervirens L.) Clones 52 and 30 under controlled conditions of high irradiation (about 1900 micromol m(-2) s(-1) for 60 min; HL treatment), followed by 60 min in darkness. The degree of photoinhibition was determined based on the ratio of variable to maximum chlorophyll fluorescence (Fv/Fm), which is a measure of the potential efficiency of photosystem II (PSII), and on electron transport measurements. The Fv/Fm ratio declined in needles of both clones in response to the HL treatment. Minimal fluorescence (Fo) increased in HL-treated needles of both clones. The HL treatment decreased rates of whole-chain and PSII activity of isolated thylakoids more in Clone 52 than in Clone 30. In needles of both clones, PSI activity was less sensitive to photoinhibition than PSII activity. In the subsequent 60-min dark incubation, fast recovery was observed in needles of both clones, with PSII efficiencies reaching similar values to those in non-photoinhibited needles. The artificial exogenous electron donors diphenyl carbazide (DPC), hydroxylamine (NH2OH) and manganese chloride (MnCl2) failed to restore the HL-induced loss of PSII activity in needles of Clone 30, whereas DPC and NH2OH significantly restored PSII activity in photoinhibited needles of Clone 52. Quantification of the PSII reaction center protein D1 and the 33-kDa protein of the water-splitting complex following HL treatment of needles revealed pronounced differences between Clone 52 and Clone 30. The large decrease in PSII activity in HL-treated needles was caused by the marked loss of D1 protein and 33-kDa protein in Clone 30 and Clone 52, respectively.  相似文献   

15.
Effects of winter snow cover on photoinhibition and possible interactions with winter desiccation were investigated in situ in an evergreen subalpine woody species, Rhododendron ferrugineum L., at the alpine timberline (1950 m a.s.l.). Timing and duration of complete snow cover markedly influenced potential efficiency of photosystem II (PSII; F(v) /F(m)). Lack of snow cover led to severe but mostly reversible photoinhibition with F(v)/F(m) values as low as 0.05. Complete snow cover immediately stopped further reductions in PSII efficiency. Snow cover promoted recovery from photoinhibition, but only if, in addition to shading by snow, plants were exposed to nonfreezing temperatures close to 0 degrees C. The F(v)/F(m) ratio was closely related to minimum leaf temperatures because both photoinhibition and recovery from photoinhibition were strongly influenced by temperature. The period without major reductions in PSII efficiency lasted for only two months, reflecting the extremely short growing period in the subalpine environment. Compared with complete snow cover, incomplete snow cover led to significantly higher water losses as well as lower dehydration tolerance, because both osmotic adjustment and changes in turgor maintenance capacity were significantly reduced. Interactions between photoinhibition and winter desiccation were masked by the direct effects of freezing temperatures. However, both photoinhibition and winter desiccation are closely linked and occur together under field conditions in this evergreen subalpine woody species.  相似文献   

16.
The ecophysiological mechanisms underlying plant-plant interactions and forest regeneration processes in Mediterranean ecosystems are poorly understood, and the experimental evidence for the role of light availability in these processes is particularly scant. We analyzed the effects of high and low irradiances on 31 ecological, morphological and physiological variables in saplings of four late-successional Mediterranean trees, two deciduous (Acer opalus subsp. granatense (Boiss.) Font Quer & Rothm. and Quercus pyrenaica Willd.) and two evergreen (Pinus nigra Arnold subsp. salzmannii (Dunal) Franco and Quercus ilex L.), which coexist in mature montane forests. Species differed in both their capacity to withstand high radiation and in their shade tolerance. The two deciduous species were the least tolerant to high radiation, exhibiting both dynamic and chronic photoinhibition in full sunlight, with severe implications for gas exchange and photosynthetic performance. Excess light severely limited the survival of A. opalus subsp. granatense, even minor reductions of excessive radiation (from full sunlight to 80% sunlight) being crucial for sapling survival. Among species, P. nigra was the most tolerant of high irradiances but showed the poorest shade acclimation. Interspecific differences in the mechanisms of response to light provide a partial explanation of the differential regeneration patterns previously reported for these species, with shade-tolerant plants (i.e., deciduous broadleaf species) benefiting the most from associations with nurse plants. We conclude that light availability is an important environmental factor defining the regeneration niche of Mediterranean woody species.  相似文献   

17.
We investigated whether timing and rate of growth are related to the life strategies and fitness of three conifer species. Intra-annual dynamics of wood formation, shoot elongation and needle phenology were monitored over 3 years in five Norway spruces (Picea abies (L.) Karst.), five Scots pines (Pinus sylvestris L.) and five silver firs (Abies alba Mill.) grown intermixed. For the three species, the growing season (delimited by cambial activity onset and cessation) lasted about 4 months, while the whole process of wood formation lasted 5-6 months. Needle unfolding and shoot elongation followed the onset of cambial activity and lasted only one-third of the season. Pines exhibited an 'extensive strategy' of cambial activity, with long durations but low growth rates, while firs and spruces adopted an 'intensive strategy' with shorter durations but higher growth rates. We estimated that about 75% of the annual radial increment variability was attributable to the rate of cell production, and only 25% to its duration. Cambial activity rates culminated at the same time for the three species, whereas shoot elongation reached its maximal rate earlier in pines. Results show that species-specific life strategies are recognizable through functional traits of intra-annual growth dynamics. The opposition between Scots pine extensive strategy and silver fir and Norway spruce intensive strategy supports the theory that pioneer species are greater resource expenders and develop riskier life strategies to capture resources, while shade-tolerant species utilize resources more efficiently and develop safer life strategies. Despite different strategies, synchronicity of the maximal rates of cambial activity suggests a strong functional convergence between co-existing conifer species, resulting in head-on competition for resources.  相似文献   

18.

Key message

In tree communities, tree size inequality reduces productivity and interacts with tree shade tolerance to modulate stand productivity, with a higher productivity in stands where shade-intolerant species dominate shade-tolerant species in size.

Context

Positive diversity–productivity relationships have been reported in different plant communities, including tree communities. These effects may be strongly related to both structural diversity and functional diversity, but also to their interactions if there is a non-random distribution of species functional characteristics among canopy layers.

Aims

We explore the relative effects on forest productivity of tree species diversity, tree size inequality, and species shade tolerance diversity, as well as the effect of the distribution of tree shade tolerance in the canopy.

Methods

We used 11,054 mixed-species forest plots from the French Forest Inventory (IGN) distributed throughout France (2006–2011). We analyzed the effects of species richness, shade tolerance diversity, and height inequality on forest plot productivity, represented by basal area annual increment over a period of 5 years, while controlling for first-order structure characteristics (basal area and quadratic mean diameter) and environmental factors (soil water budget and sum of growing degree days). Using the covariance between tree height and shade tolerance in mixed species canopies, we also explored the effect of the distribution of species’ shade tolerance among canopy layers.

Results

The results showed a positive effect of species richness (effect size, 0.02) and a negative effect of height inequality (??0.05) on mixed-forest productivity. We also showed that a negative covariance between shade tolerance and height (e.g., higher proportion of shade-tolerant species in lower height classes) increased productivity (0.01). Shade tolerance diversity did not affect productivity.

Conclusion

In tree communities, as shown previously in monospecific forest stands, tree size inequality reduces productivity. This effect is modulated by the distribution of shade tolerance among canopy layers. Previous studies on species diversity effect have generally overlooked the importance of the size structure and the size hierarchy of functional characteristics. These effects are, however, crucial and deserve to be explored in greater detail.
  相似文献   

19.
We investigated survival and growth responses of planted and advance natural regeneration species of varying shade tolerance to partial retention harvesting in moist warm Interior Cedar-Hemlock (ICHmw2) and dry cool Montane Spruce (MSdk) ecosystems of southeastern British Columbia, Canada. Treatments included three levels of overstory basal area retention (none, light (∼25%), or heavy (∼50%)) installed by two harvest methods (handfelled or a pushover falling technique being tested for its ability to control the spread of root disease). After 10 years, growth of both planted and natural regeneration species of varying shade tolerance tended to increase with decreasing overstory retention and associated increases in light availability. In contrast, significant survival responses to retention level were lacking except in the case of shade-intolerant western larch. Harvest method had a variable effect on regeneration survival and growth. Where significant responses did occur, they were generally attributed to harvesting effects on the characteristics of planting microsites rather than root disease spread. Natural regeneration densities at the ICHmw2 site were high at all retention levels, whereas stocking was less consistent at the MSdk site. We concluded that moderately shade-tolerant to shade-tolerant interior spruce and western redcedar can, under conditions similar to those of our study sites, be successfully established under overstories of up to approximately 25 m2/ha basal area, but that growth performance is likely to be significantly lower than in clearcuts. Despite early survival issues, conclusions regarding Douglas-fir were similar. Poorer survival and vigour of shade-intolerant western larch suggested this species is not suitable for regeneration in partial retention systems where timber production is the primary objective. Where non-timber objectives predominate, survival and acceptable growth of even a small proportion of larch could add to the diversity of the regenerating stand.  相似文献   

20.
三种松树叶绿素荧光特征比较研究   总被引:1,自引:0,他引:1  
以赤松、油松、白皮松3种松树为材料,利用便携式调制荧光仪MINI-PAM测定自然条件下其各叶绿素荧光参数Ft、Fm、Yield、ETR、qP、qN或NPQ和PAR指标的日变化。结果表明:3种松树各个时段荧光参数值在总体上差异不大,叶绿素荧光参数的变化依赖于PAR的变化。Ft、rETR、qN和NPQ的变化与PAR的变化趋势一致,Fm、Yield和qP的变化与PAR的变化相反。其中,白皮松的量子产量Yield、相对光合电子传递速率ETR、光化学淬灭系数qP、非光化学淬灭系数qN和NPQ均高于赤松和油松,说明其能通过增加热耗散来尽量降低光抑制程度,对当地环境较为适应。本研究为3种松树抗性研究、苗木管理、科学种植等提供了生理指标和依据。  相似文献   

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