Cherrybark oak stump sprout survival and development five years following plantation thinning in the lower Mississippi alluvial valley,USA

Cherrybark oak (Quercus pagoda Raf.) stump sprouts were studied for 5 years in a 30-year-old plantation thinned to 70–75% stocking (light thinning) and 45–50% stocking (heavy thinning). Sprouting success, survival, number of sprouts per stump, and sprout height differed little between thinning treatments throughout the 5-year study period. Pre-harvest tree d.b.h. also had no influence on sprout survival and development. A 2-year drought reduced survival and may have influenced sprout development. Sprout clump survival dropped from 90% 1 year following thinning to 46% 3 years after thinning. Although sprout height averaged 337 cm 5 years after thinning, annual sprout growth decreased from 166 cm the first year after thinning to 33 cm in each of the last 2 growing seasons. Results indicated that bottomland hardwood regeneration evaluation models may underestimate the potential of oak stump sprouts to contribute to pre-harvest regeneration assessments. Further study in the role of stump sprouts to regenerate bottomland oak species is needed. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged     

Water use by Tabor and Kermes oaks growing in their respective habitats in the Lower Galilee region of Israel

We estimated water use by the two main oak species of the Lower Galilee region of Israel—Tabor (Quercus ithaburensis) and Kermes (Quercus calliprinos)—to develop management options for climate-change scenarios. The trees were studied in their typical phytosociological associations on different bedrock formations at two sites with the same climatic conditions. Using the heat-pulse method, sap flow velocity was measured in eight trunks (trees) of each species during a number of periods in 2001, 2002 and 2003. Hourly sap flux was integrated to daily transpiration per tree and up-scaled to transpiration at the forest canopy level. The annual courses of daytime transpiration rate were estimated using fitted functions, and annual totals were calculated. Sap flow velocity was higher in Tabor than in Kermes oak, and it was highest in the youngest xylem, declining with depth into the older xylem. Average daytime transpiration rate was 67.9 ± 4.9 l tree⁻¹ d⁻¹, or 0.95 ± 0.07 mm d⁻¹, for Tabor oak, and 22.0 ± 1.7 l tree⁻¹d⁻¹, or 0.73 ± 0.05 mm d⁻¹, for Kermes oak. Differences between the two oak species in their forest canopy transpiration rates occurred mainly between the end of April and the beginning of October. Annual daytime transpiration was estimated to be 244 mm year⁻¹ for Tabor oak and 213 mm year⁻¹ for Kermes oak. Adding nocturnal water fluxes, estimated to be 20% of the daytime transpiration, resulted in total annual transpiration of 293 and 256 mm year⁻¹ by Tabor and Kermes oaks, respectively. These amounts constituted 51% and 44%, respectively, of the 578 mm year⁻¹ average annual rainfall in the region. The two species differed in their root morphology. Tabor oak roots did not penetrate the bedrock but were concentrated along the soil–rock interface within soil pockets. In contrast, the root system of Kermes oak grew deeper via fissures and crevices in the bedrock system and achieved direct contact with the deeper bedrock layers. Despite differences between the two sites in soil–bedrock lithological properties, and differences in the woody structure, annual water use by the two forest types was fairly similar. Because stocking density of the Tabor oak forests is strongly related to bedrock characteristics, thinning as a management tool will not change partitioning of the rainfall between different soil pockets, and hence soil water availability to the trees. In contrast, thinning of Kermes oak forests is expected to raise water availability to the remaining trees.     

Survival and development of Liaodong oak stump sprouts in the Huanglong Mountains of China six years after three partial harvests

The survival and development of Liaodong oak (Quercus liaotungensis Koidz) stump sprouts were studied for 6 years under leave-tree, as well as 60 and 75% crown closure uniform shelterwood in the Huanglong Mountains of China. Stump sprout survival rate, number of sprouts, and sprout height under the three partial harvests during the first year post-harvest did not differ. Six years post-harvest, 78.32% of the stumps had living sprouts. No variation in sprout survival was found among the treatments throughout the study period. The number of living sprouts per stump was 13.77 ± 0.50 in the first growing season after harvest, and dropped to 6.49 ± 0.22 after 6 years. This result indicated the occurrence of self-thinning within the sprout clumps. Moreover, the height of dominant sprout increased from 52.88 ± 1.62 cm (in the first year) to 132.82 ± 2.35 cm (in the sixth year) post-harvest. Sprout height in the leave-tree harvest type was higher than those in the two shelterwoods in the fifth and sixth years after logging. Annual height growth was approximately 8–96 cm during the first growing season, and then decreased at a rate of 1–44 cm per year in the following years. A negative relationship was found between sprout height and the number of sprouts per stump. This result indicated a trade-off between the number of sprouts and height growth. The number of sprouts per stump decreased by about 0.2 stem for each 1 m²/ha increased in residual basal area. Therefore, sprouting may play an important role in the restoration of the secondary forest after harvesting.     

Development of water oak stump sprouts under a partial overstory

A 28-year-old water oak (Quercus nigra L.) plantation was thinned from below to either 254 or 462 stems per hectare to determine the influence of a partial canopy on oak stump sprout development. Sprout clump survival, number of living sprouts in a clump, and height and DBH of the dominant sprout in a clump were measured in years 1–5 and 7 after harvest. By year 7, sprout clump survival under the heavily thinned canopy was 23% higher than under the lightly thinned canopy. Within-clump sprout mortality was not influenced by overstory thinning level, but by year 2 height and diameter increment were 15% and 22% greater, respectively, under the heavy overstory removal. Positive height and diameter growth of dominant sprouts continued under both canopy conditions through year 7, but early benefits of the heavy overstory removal on sprout growth diminished. Additional overstory removal or sprout clump thinning would be necessary to sustain sprout clump survival and sprout growth over extended periods.     

Growth and nutrition of Pinus radiata in response to fertilizer applied after thinning and interaction with defoliation associated with Essigella californica

Infestations of Essigella californica following the installation of post-thinning fertilizer trials in Pinus radiata plantations provided an opportunity to examine the impact of repeated defoliation over a period of 8 years (1997–2005). Replicated treatments (n = 4) of nil fertilizer (control), N (300 kg ha⁻¹) as urea, P (80 kg ha⁻¹) and S (45 kg ha⁻¹) as superphosphates were applied immediately after thinning at three sites and this was followed by a second application of NPS fertilizers 6 years later with N applied at 300 kg ha⁻¹ as urea and ammonium sulphate and P at 80 or 120 kg ha⁻¹. Defoliation of untreated P. radiata gradually increased to 50% over a period of 8 years. Basal area growth was negatively correlated with average defoliation for two consecutive post-fertilizer periods of 6 and 2 years. Growth responses to fertilizer varied considerably between sites but the largest improvement in growth was due to NPS fertilizer, this increased basal area by 30–80%. Application of N fertilizer raised total N levels in foliage and increased defoliation with a commensurate loss in growth under conditions of deficiencies of S or P. Repeated infestations gradually increased the percentage of trees with severe defoliation (>80% loss of foliage) indicating that nutrient-deficient trees have a reduced capacity for foliage recovery between episodes of peak infestation. In contrast, treatment with N fertilizer in combination with S- and P-corrected deficiencies of these nutrients, raised levels of total N in foliage and reduced defoliation to approximately 20%. Basal area growth responses to NPS fertilizers reflected improved nutrition as well as reduced insect damage. The reduction in defoliation under conditions of balanced tree nutrition was most likely due to enhanced needle retention following correction of P deficiency as well as greater availability of nutrients enabling a more vigorous recovery of P. radiata after an episode of E. californica activity. Treatment with fertilizer therefore reduced the long-term impact of aphid damage and improved growth of P. radiata.     

辽东栎伐桩萌苗的发育规律

通过对黄土高原黄龙山林区辽东栎林固定样地连续6年的调查,研究间伐强度分别为13.4％(弱度间伐)和30.0％(强度间伐)的辽东栎伐桩萌苗的存活与生长规律.结果表明:1)间伐第1年,辽东栎伐桩均有萌生枝发生,每桩萌生枝数量和萌生枝高度在不同强度间无显著差异;2)萌生枝数量受伐桩高度和直径的影响,当伐桩高度较低时,萌生枝数量随伐桩高度的增加而增加,超过20cm后随之减少,呈三次抛物线关系;伐桩萌生枝数量随着伐桩直径增加呈现先增加后减少的趋势,直径30～50cm的伐桩萌生枝数量最多;3)伐桩萌生枝数量与最长萌生枝高度呈显著线性负相关,存在“萌生枝数量——高度权衡”现象;4)间伐第6年,伐桩的存活率为97.01％,强度间伐区萌生枝高度显著高于弱度间伐区;5)调查期间,辽东栎伐桩萌生枝数量逐年下降,伐桩产生的萌生枝数量越多,后期萌生枝死亡的数量也越多,表明伐桩能够自我稀疏,存在“萌代主”现象.在黄土高原地区,辽东栎伐桩萌芽潜力很大,应合理利用;间伐时伐桩高度应尽量低于20cm;伐桩直径在30～50cm更有利于其无性更新;间伐后伐桩应及时除蘖定株,促进萌生枝生长;间伐强度30.0％(强度间伐)更有利于辽东栎伐桩萌生枝的生长.     

Thinning method and intensity influence long-term mortality trends in a red pine forest

Tree mortality shapes forest development, but rising mortality can represent lost production or an adverse response to changing environmental conditions. Thinning represents a strategy for reducing mortality rates, but different thinning techniques and intensities could have varying impacts depending on how they alter stand structure. We analyzed trends in stand structure, relative density, stand-scale mortality, climate, and correlations between mortality and climate over 46 years of thinning treatments in a red pine forest in Northern Minnesota, USA to examine how thinning techniques that remove trees of different crown classes interact with growing stock manipulation to impact patterns of tree mortality. Relative density in unharvested plots increased during the first 25 years of the study to around 80%, then began to plateau, but was lower (12–62%) in thinned stands. Mortality in unharvested plots claimed 2.5 times more stems yr⁻¹ and 8.6 times as large a proportion of annual biomass increment during the last 21 years of the study compared to the first 25 years, but showed few temporal trends in thinned stands. Mortality in thinning treatments was generally lower than in controls, particularly during the last 21 years of the study when mortality averaged about 0.1% of stems yr⁻¹ and 4% of biomass increment across thinning treatments, but 0.8% of stems yr⁻¹ and 49% of biomass increment in unharvested plots. Treatments that combined thinning from above with low growing stock levels represented an exception, where mortality exceeded biomass production after initial thinning. Mortality averaged less than 0.1% of stems yr⁻¹ and less than 1% of annual biomass production in stands thinned from below. These trends suggest thinning from below minimizes mortality across a wide range of growing stock levels while thinning from above to low growing stock levels can result in dramatic short-term increases in mortality. Moderate to high growing stock levels (21–34 m² ha⁻¹) may offer greater flexibility for limiting mortality across a range of thinning methods. Mean and maximum annual and growing season temperatures rose by 0.6–1.8 °C during the study, and temperature variables were positively correlated with mortality in unharvested plots. Mortality increases in unharvested plots, however, were consistent with self-thinning principles and probably not driven by rising temperatures. These results suggest interactions between thinning method and intensity influence mortality reductions associated with thinning, and demonstrate the need for broader consideration of developmental processes as potential explanations for increased tree mortality rates in recent decades.     

林龄、采伐方式对大叶相思萌芽更新的影响*

在海南省琼海县进行了不同林龄、不同采伐方式对大叶相思萌芽更新影响的研究，同时比较了保留不同萌条数量的萌芽林生长表现。结果表明：（１）林龄对伐桩萌芽率没有显著影响，但对伐桩存活率及萌条的径、高生长影响显著或极显著。林龄越大，萌芽更新效果越差；（２）皆伐和隔行采伐对大叶相思伐桩的萌芽率、萌条数量和存活率均无显著影响，但对萌条的径、高生长影响极显著。试验证明对大叶相思林分施以隔行采伐通过萌芽更新建立复层     

Analysis of nutrient depletion in a radiata pine plantation

A trial in an 11-year-old stand of radiata pine (Pinus radiata D. Don) was used to analyse the effects of accelerated loss of nutrients from the site on forest productivity and nutrient status. Raking of litter was undertaken over 14 years prior to thinning, then for 2 years after thinning at which time the trial was destroyed in a wind storm. The experimental design was a factorial of three main treatments: (i) removal (raking) versus nil removal of the forest floor, (ii) replacement or non replacement of nutrients to adjust for imbalances between nutrients in litter and those in the tree stem, and (iii) complete replacement (or not) of all nutrients removed in the litter. Additionally, a small trial was incorporated to address components of physical aspects of litter removal by comparing raking with ‘raking and a cover of woven plastic mesh’. Raking and nutrient additions were carried out approximately every 6 months.Over the study period, the raking treatment removed about 75 Mg ha⁻¹ of organic material with contained nutrients (559 kg ha⁻¹ of N, 68 kg ha⁻¹ of P, 323 kg ha⁻¹ of Ca, 91 kg ha⁻¹ of Mg, 243 kg ha⁻¹ of K, 0.9 kg ha⁻¹ of B) and this related to about four normal sawlog harvests or one total tree harvest. Up to the time of thinning, raking reduced basal area increment by 25% while raking together with replacement of nutrients reduced this by about 12%. Nutrient additions to unraked plots led to increases of up to 14% in basal area increment. The raking treatment reduced foliage nitrogen and this was correlated with reduced growth while other nutrients such as boron and sulphur were reduced but not to a degree to affect growth or health. The results were used to assess the effects on soil nutrient status and growth of different harvesting regimes (wood only, wood plus bark, total tree).     

Aboveground biomass and nutrient accumulation dynamics in young black alder, silver birch and Scots pine plantations on reclaimed oil shale mining areas in Estonia

The growth, aboveground biomass production and nutrient accumulation in black alder (Alnus glutinosa (L.) Gaertn.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) plantations during 7 years after planting were investigated on reclaimed oil shale mining areas in Northeast Estonia with the aim to assess the suitability of the studied species for the reclamation of post-mining areas. The present study revealed changes in soil properties with increasing stand age. Soil pH and P concentration decreased and soil N concentration increased with stand age. The largest height and diameter of trees, aboveground biomass and current annual production occurred in the black alder stands. In the 7-year-old stands the aboveground biomass of black alder (2100 trees ha⁻¹) was 2563 kg ha⁻¹, in silver birch (1017 trees ha⁻¹) and Scots pine (3042 trees ha⁻¹) stands respective figures were 161 and 1899 kg ha⁻¹. The largest amounts of N, P, K accumulated in the aboveground part were in black alder stands. In the 7th year, the amount of N accumulated in the aboveground biomass of black alder stand was 36.1 kg ha⁻¹, the amounts of P and K were 3.0 and 8.8 kg ha⁻¹, respectively. The larger amounts of nutrients in black alder plantations are related to the larger biomass of stands. The studied species used N and P with different efficiency for the production of a unit of biomass. Black alder and silver birch needed more N and P for biomass production, and Scots pine used nutrients most efficiently. The present study showed that during 7 years after planting, the survival and productivity of black alder were high. Therefore black alder is a promising tree species for the reclamation of oil shale post-mining areas.     

Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Soil changes induced by Acacia mangium plantation establishment: Comparison with secondary forest and Imperata cylindrica grassland soils in South Sumatra,Indonesia

Acacia plantation establishment might cause soil acidification in strongly weathered soils in the wet tropics because the base cations in the soil are translocated rapidly to plant biomass during Acacia growth. We examined whether soils under an Acacia plantation were acidified, as well as the factors causing soil acidification. We compared soils from 10 stands of 8-year-old Acacia mangium plantations with soils from 10 secondary forests and eight Imperata cylindrica grasslands, which were transformed into Acacia plantations. Soil samples were collected every 5–30 cm in depth, and pH and related soil properties were analyzed. Soil pH was significantly lower in Acacia plantations and secondary forests than in Imperata grasslands at every soil depth. The difference was about 1.0 pH unit at 0–5 cm and 0.5 pH unit at 25–30 cm. A significant positive correlation between pH and base saturation at 0–20 cm depth indicated that the low pH under forest vegetation was associated with exchangeable cation status. Using analysis of covariance (ANCOVA), with clay content as the covariate, exchangeable Ca (Ex-Ca) and Mg (Ex-Mg) stocks were significantly lower in forested areas than in Imperata grasslands at any clay content which was strongly related to exchangeable cation stock. The adjusted average Ex-Ca stock calculated by ANCOVA was 249 kg ha⁻¹ in Acacia plantations, 200 kg ha⁻¹ in secondary forests, and 756 kg ha⁻¹ in Imperata grasslands at 0–30 cm. Based on a comparison of estimated nutrient stocks in biomass and soil among the vegetation types, the translocation of base cations from soil to plant biomass might cause a decrease in exchangeable cations and soil acidification in Acacia plantations.     

The effect of land use type on net nitrogen mineralization on abandoned agricultural land: Silver birch stand versus grassland

The effect of land use type on the dynamics and annual rate of net nitrogen mineralization (NNM) in a naturally generated silver birch stand and in a grassland, both on abandoned agricultural land, was assessed in situ in the upper 0–20 cm soil layer using the method of buried polyethylene bags. Annual NNM rate in the birch stand (156 kg N ha⁻¹ year⁻¹) was higher than in the grassland (102 kg N ha⁻¹ year⁻¹); in both cases NNM covered a major part of the plants annual nitrogen demand. The rate of NNM in the upper 0–10 cm soil layer in the birch stand (99 kg N ha⁻¹ year⁻¹) exceeded the respective rate of NNM in the grassland (51 kg N ha⁻¹ year⁻¹) roughly two times. In the grassland the rates of NNM in the 0–10 and 10–20 cm layers were equal; in the birch stand NNM in the 0–10 cm layer was 1.7 times higher than in deeper 10–20 cm layer. The intensity of daily NNM in the upper 0–10 cm soil layer in the birch stand was the highest in June and in the grassland in May, 776 and 528 mg kg⁻¹ N day⁻¹, respectively. In our study no significant correlation was found between NNM and the environmental factors monthly mean soil temperature, soil moisture content and pH.     

Early regeneration of Fraxinus rhynchophylla in the understorey of Larix kaempferi stands in response to thinning

The effects of thinning were investigated on naturally regeneratedFraxinus rhynchophylla Hance (Korean ash) grown in the understoreyof a Larix kaempferi (Lamb.) Carr. plantation, established ona site that previously supported old growth hardwood forests.The regeneration characteristics were evaluated after thinningof the plantation (about 60 per cent of relative density wasremoved). In the naturally regenerated and mixed stands, Quercusmongolica Fisch. was the most abundant tree species in the overstorey,but the understorey of the thinned plantations was dominatedby F. rhynchophylla with six other minor tree species. Fraxinusrhynchophylla was regenerated by four propagation types includingnatural seedling, stump sprout, root sprout and layering. Beforethinning, the difference of average height growth for each regenerationtype was insignificant. One year after thinning, the stump sproutsshowed the fastest growth among the regeneration types. Heightgrowth of advance regeneration was slow in the year followingthinning but then increased rapidly surpassing the other regenerationtypes 5 years after thinning. Thinning had an effect on thecomposition of regenerated trees also by seedling, stump sprout,root sprout and layering in the understorey. Advance regenerationsoccupied the upper understorey at 5 years after thinning. Alarge number of stump sprouts emerged in the plots 1 year afterthinning, while at 5 years after thinning the occupation ofstump sprouts decreased. Root sprouts and layers faded away.Considering the early growth and the composition of understoreyF. rhynchophylla, its advance regeneration has a high potentialto develop into overstorey trees at this site.     

Nitrogen leaching in response to increased nitrogen inputs in subtropical monsoon forests in southern China

Dissolved inorganic nitrogen (DIN) (as ammonium nitrate) was applied monthly onto the forest floor of one old-growth forest (>400 years old, at levels of 50, 100 and 150 kg N ha⁻¹ yr⁻¹) and two young forests (both about 70 years old, at levels of 50 and 100 kg N ha⁻¹ yr⁻¹) over 3 years (2004–2006), to investigate how nitrogen (N) input influenced N leaching output, and if there were differences in N retention between the old-growth and the young forests in the subtropical monsoon region of southern China. The ambient throughfall inputs were 23–27 kg N ha⁻¹ yr⁻¹ in the young forests and 29–35 kg N ha⁻¹ yr⁻¹ in the old-growth forest. In the control plots without experimental N addition, a net N retention was observed in the young forests (on average 6–11 kg N ha⁻¹ yr⁻¹), but a net N loss occurred in the old-growth forest (−13 kg N ha⁻¹ yr⁻¹). Experimental N addition immediately increased DIN leaching in all three forests, with 25–66% of added N leached over the 3-year experiment. At the lowest level of N addition (50 kg N ha⁻¹ yr⁻¹), the percentage N loss was higher in the old-growth forest (66% of added N) than in the two young forests (38% and 26%). However, at higher levels of N addition (100 and 150 kg N ha⁻¹ yr⁻¹), the old-growth forest exhibited similar N losses (25–43%) to those in the young forests (28–43%). These results indicate that N retention is largely determined by the forest successional stages and the levels of N addition. Compared to most temperate forests studied in Europe and North America, N leaching loss in these seasonal monsoon subtropical forests occurred mainly in the rainy growing season, with measured N loss in leaching substantially higher under both ambient deposition and experimental N additions.     

Potential long-term growth gains from early clump thinning of coppice-regenerated oak stands

A study was established in the Missouri Ozarks to evaluate coppice regeneration of oak. Five years after a 32-year-old stand was clearcut and regenerated naturally through stump sprouts, the dominant sprout on each stump was identified based upon height. Treated plots were thinned to the single dominant sprout on each stump whereas control plots were not thinned. Twenty-five years later the largest 247, 371, 494, and 618 stems per ha were examined and height of the dominant sprout at age 5 was found to be strongly related to dbh at age 30 in both thinned and unthinned plots. However, in the thinned plots, the largest 494 stems per ha were on average 11%, 28%, and 58% greater, respectively, in height, dbh, and volume compared to similar dominant sprouts in unthinned plots. Logistic regression analysis was used to develop curves for the evaluation of potential gains from clump thinning. In this analysis, the average height of a stand at age 5 was used to estimate thinning gains at age 30.     

Estimating biomass production and carbon storage for a fast-growing makino bamboo (Phyllostachys makinoi) plant based on the diameter distribution model

The purpose of this study was to estimate biomass and carbon storage for a fast-growing makino bamboo (Phyllostachys makinoi). The study site was located in central Taiwan and the makino bamboo plantation had a stand density of 21191 ± 4107 culms ha⁻¹. A diameter distribution model based on the Weibull distribution function and an allometric model was used to predict aboveground biomass and carbon storage. For an accurate estimation of carbon storage, the percent carbon content (PCC) in different sections of bamboo was determined by an elemental analyzer. The results showed that bamboos of all ages shared a similar trend, where culms displayed a carbon storage of 47.49–47.82%, branches 45.66–46.23%, and foliage 38.12–44.78%. In spite of the high density of the stand, the diameter distribution of makino bamboo approached a normal distribution and aboveground biomass and carbon storage were 105.33 and 49.81 Mg ha⁻¹, respectively. Moreover, one-fifth of older culms from the entire stand were removed by selective cutting. If the distribution of the yield of older culms per year was similar to the current stand, the yields of biomass and carbon per year would be 21.07 and 9.89 Mg ha⁻¹ year⁻¹. An astonishing productivity was observed, where every 5 years the yield of biomass and carbon was equal to the current status of stockings. Thus, makino bamboo has a high potential as a species used for carbon storage.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.