Seasonal and spatial variations in leaf nitrogen content and resorption in a Quercus serrata canopy

To elucidate the relationships between spatiotemporal changes in leaf nitrogen (N) content and canopy dynamics, changes in leaf N and distribution in the canopy of a 26-year-old deciduous oak (Quercus serrata Thunb. ex. Murray) stand were monitored throughout the developmental sequence from leaf expansion to senescence, by estimating the leaf mass and N concentrations of all the canopy layers. Seasonal changes were observed in leaf N concentration per unit leaf dry mass (N (m)), which peaked after bud burst, declined for two weeks shortly thereafter, and then remained constant for the rest of the growing season for each canopy layer. Leaf N concentration per unit leaf area (N (a)) was higher in the upper layer than in the lower layer throughout the growing season, and was closely correlated with relative irradiance (RI) in the summer when the air temperature was moderately high. The N concentrations of all leaf layers started to decrease in November, and reached their lowest values in late November, whereas LMA scarcely changed throughout the season. The lowest N concentrations did not differ significantly among the canopy layers. Seasonal changes in the relationship between N (a) and RI were detected, indicating that N (a) is optimized temporally as well as spatially. Nitrogen resorption efficiency was highest in the upper canopy layers where larger amounts of N were invested. Based on the estimates of leaf mass and leaf N concentrations of the canopy layers, total leaf N concentration of the whole canopy was estimated to be 84.1 kg ha(-1) in the summer, and 37.3 kg ha(-1) in late November. Therefore, 46.8 kg ha(-1) of leaf N in the canopy (about 56% of the total N) was resorbed just before leaf abscission.     

Relative shoot height and irradiance and the shoot and leaf properties of Quercus serrata saplings

Effects of relative shoot height and irradiance on shoot and leaf properties of Quercus serrata Thunb. saplings growing in the understory and in gaps were investigated. Photosynthetic photon flux (PPF) at the location of the shoot relative to that in the open (relative PPF; rPPF) and the height of the shoot base relative to tree height (relative height; rHeight) were measured for all current-year shoots of each sapling. Current-year shoot properties (length, leaf area, number of daughter shoots) and mortality, and leaf properties (mass per area (LMA) and nitrogen content per area (N(area))) were examined in relation to rPPF and rHeight. N(area) was used as a proxy for area-based assimilative capacity. Shoot length, leaf area per shoot and number of daughter shoots increased with increasing rHeight, especially in well-lit conditions. Shoot mortality decreased with increasing rHeight and rPPF. Both LMA and N(area) were positively correlated with rPPF, but not rHeight.     

Virulence of Raffaelea quercivora isolates inoculated into Quercus serrata logs and Q. crispula saplings

This study was conducted to compare the virulences of various isolates of Raffaelea quercivora—a fungus that causes Japanese oak wilt disease—towards Japanese oak trees. Five isolates were collected from a wide range of Japan and inoculated into Quercus serrata logs and Q. crispula saplings. The tangential length of the discolored sapwood in the Q. serrata logs differed significantly among the isolates. The trend in isolate virulence was similar for the Q. serrata logs and the Q. crispula saplings. This is the first report suggesting that there is variability in virulence among isolates of R. quercivora.     

Variation in herbivory-induced responses within successively flushing Quercus serrata seedlings under different nutrient conditions

Herbivore damage can induce the host plant to alter the chemical and physical qualities of its leaves, which is thought to be a plant strategy—termed “induced response”—for avoiding further herbivory. In woody plants, many studies have considered variation in induced response with resource availability, but few studies have examined this variation in relation to growth patterns of woody plants. We studied the phenotypic variability of induced response within successively flushing Quercus serrata seedlings. Q. serrata seedlings were grown under controlled conditions. The controlled factors were herbivore damage (herbivore-damaged and -undamaged) and soil fertility (low and high). At each flush stage, the concentrations of condensed tannin (CT), total phenolics (TP), and nitrogen (N) in leaves were analyzed, and the leaf mass per area (LMA) was measured. CT and TP concentrations in leaves and LMA were higher in herbivore-damaged seedlings. Leaves of the first flushes showed greater sensitivity to herbivore damage and had a higher CT concentration than leaves of the later flushes. Furthermore, seedlings growing in low-fertility soil showed a greater induced response. The results suggest that the induced response of Q. serrata seedlings was related to the contributions of the tissue to current productivity. Leaves of the first flush showed a greater induced response, possibly because they play an important role in subsequent growth. The potential of Q. serrata seedlings to adjust the properties of leaves depending on herbivory and soil fertility in relation to growth patterns may be advantageous on the forest floor, where seedlings grow in soil of heterogeneous fertility and are constantly exposed to herbivory.     

Nutrient retranslocation within the foliage of Pinus sylvestris

During the period 1983-1987, retranslocation of N, P, K, Ca, Mg, Mn, Zn, Fe, B and Al within the foliage of Scots pine (Pinus sylvestris L.) was studied in sapling, pole-stage and mature stands in eastern Finland. Needle concentrations of the mobile nutrients N, P, and K varied seasonally because of retranslocation. In unfertilized plots, needle contents of N, P and K decreased 62-92% during senescence and needle dry weight decreased 19-51%. In some years, needle contents of Ca, Mn, Zn, Fe, B and Al increased during senescence. Retranslocation was more efficient from needles with greater dry weight. In the fertilized, pole-stage stand, needle dry weight did not decrease at senescence four years after fertilization and nutrient content decreased less than in the other years. In the mature stand, there were no clear differences in needle dry weight and nutrient changes during senescence between years or between unfertilized and fertilized plots.     

Acclimation of shade-developed leaves on saplings exposed to late-season canopy gaps

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.     

Optimality and nitrogen allocation in a tree canopy

Physical and functional properties of foliage were measured at a variety of microsites in a broad-leaved Nothofagus fusca (Hook. f.) ?rst. canopy. The light climate of the foliage at these sites was monitored for 39 days in the late spring and early summer with in situ sensors. Foliage nitrogen content (N), mean leaf angle, and gas exchange characteristics were all correlated with the amount of light reaching the microsites during foliage development. Foliage N content on a leaf area basis ranged between ~1 and 2.5 g N m(-2) and was highest at the brightest sites. Light-saturated photosynthetic rates ranged between ~4 and 9 micro mol m(-2) s(-1), increasing from the darkest to brightest sites. A biochemical model of photosynthesis was fitted to foliage characteristics at the different microsites and used to integrate foliage assimilation among the sites over 39 days. The actual arrangement of foliage physiological characteristics in the observed microsites led to higher total canopy rates of net assimilation than > 99% of the combinations of observed foliage characteristics randomly assigned to the observed microsites. Additional simulations first related the maximum rates of electron transport (J(max)), ribulose bisphosphate turnover (V(c,max)), and dark respiration (R(d)) of Nothofagus fusca foliage to nitrogen content and then allowed foliage N (and consequently leaf gas exchange characteristics) to vary across the canopy. The observed N allocation pattern results in greater total canopy assimilation than uniform or > 99% of the simulations with random distributions of N among the microsites (constrained so that the total N allocated was equivalent to that observed in the microsites). However, the observed pattern of N allocation places less N in the brightest microsites and results in substantially less total assimilation than a simulated canopy in which N was allocated in an optimal manner where the N distribution is such that the partial derivative of leaf assimilation (A) with respect to leaf nitrogen content, partial differential A/ partial differential N, is constant among microsites. These results suggest that other factors such as wind or herbivory reduce the integrated assimilation of high-N foliage relatively more than lower-N foliage and that a partial differential A/ partial differential N optimality criteria based only on formulations of leaf gas exchange overestimate canopy assimilation.     

Photosynthetic capacity in relation to nitrogen in the canopy of a Quercus robur, Fraxinus angustifolia and Tilia cordata flood plain forest

We measured gas exchange and various leaf parameters of ash (Fraxinus angustifolia Vahl.) and oak (Quercus robur L.) in the high canopy and of lime (Tilia cordata Mill.) in the lower canopy of a planted, 120-year-old floodplain forest in southern Moravia, Czech Republic. The high-canopy leaves of F. angustifolia and Q. robur had nitrogen concentrations on a leaf area basis (N(area)) that were twice those of low-canopy leaves of T. cordata. Upper-canopy leaves of F. angustifolia had a photosynthetic rate at light saturation (A(max)) of about 16 micromol CO2 m(-2) s(-1), whereas A(max) of the upper-canopy foliage of Q. robur achieved only about two thirds of this value. Contrary to previous investigations of photosynthetic performance in monospecific stands, leaves of the uppermost branches of T. cordata at 15-m height had the highest A(max) and transpiration rate among the species studied. Water-use efficiency (WUE) was low in T. cordata at 15-m canopy height, whereas WUE was significantly higher for Q. robur leaves at 27-m height than for the other species. Leaves of T. cordata at 15-m height showed the strongest relationship between A(max) and N(area) (R2 = 0.90) followed by F. angustifolia (R2 = 0.69). The strong correlation between photosynthesis and nitrogen concentration in T. cordata at 15 m, together with the steep regression slope for the A(max):N(area) relationship, indicated that nitrogen allocation to the photosynthetic apparatus resulted in high nitrogen-use efficiency of light-saturated photosynthesis (PNUE). Despite differences in PNUE among species, PNUE was fairly constant for leaves sampled from the same canopy position, suggesting that single-leaf parameters are matched to optimize PNUE for prevailing light conditions. High PNUE in T. cordata at 15 m partially compensated for the species' subordinate position in the canopy, and may be an important mechanism for its coexistence in highly structured vegetation.     

Effects of partial defoliation on carbon and nitrogen partitioning and photosynthetic carbon uptake by two-year-old cork oak (Quercus suber) saplings

At the end of the growing season in late July, 20-month-old cork oak (Quercus suber L.) saplings were partially defoliated (63% of leaf area) to evaluate their ability to recover leaf area after defoliation. At 18 and 127 days after defoliation, changes in starch and nitrogen pools were determined in leaves and perennial organs, and variations in photosynthetic carbon uptake were investigated. To determine the role of stored nitrogen in regrowth after defoliation, plant nitrogen was labeled in the previous winter by enriching the nutrient solution with 15N. Plants recovered the lost leaf area in 127 days. Although there was remobilization of starch and nitrogen from leaves and perennial organs, the availability of resources for growth in the following spring was not decreased by defoliation. On the contrary, starch concentration in coarse roots was higher in defoliated saplings than in control saplings, presumably as a result of the higher net CO2 exchange rate in newly developed leaves compared with pre-existing leaves.     

Isolation of endophytic streptomycetes from above- and belowground organs of Quercus serrata

The occurrence and localization of endophytic actinomycetes within diverse organs of host plants provide ecological information that can be used to evaluate the significance of their spatial habitats. The aim of this study was to isolate and characterize endophytic actinomycetes in different organs of Quercus serrata. For this purpose, actinomycete isolates were obtained from surface-sterilized tissues of both symptomless seedlings and aged trees of Q. serrata and rhizosphere soil of the sampled seedlings. Thirty-five isolates with the ability to sporulate, including 4 from leaves of the aged trees, 10, 6, and 15 from leaves, stems, and roots of the seedlings, respectively, and 8 soil-derived isolates, were selected and characterized. The 16S rDNA nucleotide sequence analyses revealed that all of them belonged to the genus Streptomyces. According to a neighbor-joining phylogenetic tree constructed based on the results, the isolates of plant origin were divided into three major clades with high bootstrap values of 98 or 99 %, whereas eight soil-derived isolates were located at different positions from those of the endophyte isolates. Moreover, two larger clades were formed, one of which contained isolates derived only from aboveground parts, while the other contained isolates from all of the organs. These results suggest that the endophytic streptomycetes in Q. serrata may differ in their habitat positions (i.e., either above- or belowground parts).     

Weed control increases nitrogen retranslocation and growth of white spruce seedlings on a reclaimed oil sands soil

Early establishment of seedlings in reclaimed oil sand areas is often limited by low nutrient and water availability due to factors such as strong understory vegetation competition. Management practices such as nursery fertilization and field weed control could help early establishment of planted seedlings and reclamation success. We investigated the effect of nursery nutrient loading and field weed control on the growth, nitrogen (N) retranslocation within seedling components, and plant N uptake from the soil for white spruce (Picea glauca [Moench] Voss) seedlings planted on a highly competitive reclaimed oil sands site for two years. Exponential fertilization during nursery production increased the root biomass but not the nutrient reserve in the seedling. In the field experiment, on average across the treatments, 78 and 49% of the total N demand of new tissue growth in the first and second year were met by N retranslocation, respectively. Though exponential fertilization did not affect N retranslocation, it increased the percent height and root collar diameter growth. Weed control increased not only the growth of seedlings by increasing soil N availability, but also N retranslocation within the seedlings in the second year after outplanting. We conclude that vegetation management by weed control is feasible in improving the early growth of white spruce seedlings planted on reclaimed soils and facilitate tree establishment in the oil sands region. Optimization of the nursery exponential N fertilization regime for white spruce may further help with early revegetation of reclaimed oil sands sites.     

Throughfall,stemflow and interception loss in a mixed white oak forest (Quercus serrata Thunb.)

Throughfall and stemflow measurements in a 60-year-old white oak stand (Quercus serrata Thunb.) were carried out during two periods totalling eleven months, from August to November 1993 and from May to November 1994, in order to clarify the rainfall partitioning of this forest. Troughs and spiral-type stemflow gauges connected to tipping bucket-gauges were used for throughfall and stemflow measurements. Seventy-five storms were analyzed individually. Coefficients of variation for throughfall and stemflow ranged between 5–25% and 20–70% respectively. Partitioning of net rainfall in throughfall and stemflow represent 72% and 10% of the gross rainfall respectively. Multiple regression analyses were carried out to determine the stemflow variability. In was determined that maximum rain intensity was highly correlated with stemflow and this variable explained a further 5.5% of the stemflow variation. Estimates of averaged lag time and drainage after rain cease for stemflow were 290 and 164 min, while estimates for throughfall were 60 and 104 min. respectively. The canopy saturation was estimated from continuous storms and showed a value of 0.6 mm. The trunk storage capacity was estimated at a value of 0.2 mm. The interception loss from the forest canopy was estimated in 18%. Interception loss was heighly correlated with rainfall characteristics such as duration and intensity.     

Modeling radiation transfer within the canopy of a Chinese fir plantation

Calculation of radiation transfer within the canopy of a plantation on sloped sites is described. The canopy was assumed to consist of evenly cone-shaped crowns. Within-tree shading by object tree and between-tree shading by tree crowns around the object tree were differentiated. The two-dimensional needle area density (NAD) was introduced in the calculation of radiation transfer and interception. The model was validated with measurements of both daily photosynthetic photo flux density (PPFD) and daily photosynthetic radiation at tens of points within the crown of a 17-year-old Chinese fir (Cunninghamia lanceolata) plantation growing on a sloped site. Simulation showed that modeled data closely matched measurements. Simulation errors likely result from the uneven size of tree crowns, irregular crown shape, grouping of needles to shoots and twigs, etc. Scattering radiation amounted to below 10%, in most cases, of daily radiation, and its uncertainty was relatively small.     

Shoot structure and growth along a vertical profile within a Populus-Tilia canopy

We investigated shoot growth patterns and their relationship to the canopy radiation environment and the distribution of leaf photosynthetic production in a 27-m-tall stand of light-demanding Populus tremula L. and shade-tolerant Tilia cordata Mill. The species formed two distinct layers in the leaf canopy and showed different responses in branch architecture to the canopy light gradient. In P. tremula, shoot bifurcation decreased rapidly with decreasing light, and leaf display allowed capture of multidirectional light. In contrast, leaf display in T. cordata was limited to efficient interception of unidirectional light, and shoot growth and branching pattern facilitated relatively rapid expansion into potentially unoccupied space even in the low light of the lower canopy. At the canopy level, T. cordata had higher photosynthetic light-use efficiency than P. tremula, whereas P. tremula had higher nitrogen-use efficiency than T. cordata. However, at the individual leaf level, both species had similar efficiencies under comparable light conditions. Production of new leaf area in the canopy followed the pattern of photosynthetic production. However, the species differed substantially in extension growth and space-filling strategy. Light-demanding P. tremula expanded into new space with a few long shoots, with shoot length strongly dependent on photosynthetic photon flux density (PPFD). Production of new leaf area and extension growth were largely uncoupled in this species because short shoots, which do not contribute to extension growth, produced many new leaves. Thus, in P. tremula, the growth pattern was strongly directed toward the top of the canopy. In contrast, in shade-tolerant T. cordata, shoot growth was weakly related to PPFD and more was invested in long shoot growth on a leaf area basis compared with P. tremula. However, this extension growth was not directed and may serve as a passive means of avoiding self-shading. This study supports the hypothesis that, for a particular species, allocation patterns and crown architecture contribute as much to shade tolerance as leaf-level photosynthetic acclimation.     

侧柏栎类混交林林冠截留模拟研究

林冠截留能力是反映森林与水关系的一个重要指标,该研究充分考虑林冠结构特征与降雨特性,采用机械布样的方法,对侧柏栎类混交林林分内的3种主要水源保护林树种生长季的降雨再分配特点进行了研究。结果表明:3种树种生长季场降雨的林冠截留量都表现出随降雨量的增大而增大,截留率随降雨量的增大而减少,在降雨初期下降幅度最大,以后趋于平稳。干流量在低雨量级时随降雨量的增加而增大,但其增加幅度有限,干流率起初也均随林外降雨量的增加而呈增加的趋势。透雨量随林外降雨量的增大而增大,透雨率逐渐增大,但当林外降雨量增大到一定时透雨率基本不变。3个树种的截留量、干流量、穿透降雨量林外降雨量均呈线性相关。该文采用谢春华等的林冠截留模型对3种水源林树种进行模拟,模拟误差为11%左右,基本上可以满足林冠截留模拟需要。     

Development of Quercus robur advance regeneration following canopy reduction in an oak woodland

Observations to follow the development of 2-year-old oak seedlingstook place over a 6-year period at a fenced site within a maturestand of oak in southern England. Bramble and bracken were abundant,but the cover of other species was low. Although seedlings doubledin height, only 13 per cent survived and few were taller thaneither bramble or bracken. The initial number of seedlings wasinversely related to both canopy cover and distance to the nearestparent tree. The final number of seedlings could be predictedfrom the initial percentage covers of bramble and bracken, andeither the overstorey canopy cover or the initial number ofseedlings. The average annual rate of seedling mortality, whichwas 25–50 per cent, was significantly influenced by thevegetation present. Grass was positively related to survivalwhereas there was an inverse relationship with both brambleand bracken cover. Models estimated 80 per cent annual seedlingsurvival at 25 per cent bramble cover but only 50 per cent at60 per cent cover. Although grasses can be competitive species,results indicate that oak seedling establishment may be betterin areas with a ground flora comprising grass.     

Seasonal changes in the temperature response of photosynthesis in canopy leaves of Quercus crispula in a cool-temperate forest

Understanding seasonal changes in photosynthetic characteristics of canopy leaves is indispensable for modeling the carbon balance in forests. We studied seasonal changes in gas exchange characteristics that are related to the temperature dependence of photosynthesis in canopy leaves of Quercus crispula Blume, one of the most abundant species in cool-temperate forests in Japan. Photosynthetic rate and ribulose-1,5-bisphosphate (RuBP) carboxylation capacity (V(cmax)) at 20 degrees C increased from June to August and then decreased in September. The activation energy of V(cmax), a measure of the temperature dependence of V(cmax), was highest in summer, indicating that V(cmax) was most sensitive to leaf temperature at this time. The activation energy of V(cmax) was significantly correlated with growth temperature. Other parameters related to the temperature dependence of photosynthesis, such as intercellular CO(2) partial pressure and temperature dependence of RuBP regeneration capacity, showed no clear seasonal trend. It was suggested that leaf senescence affected the balance between carboxylation and regeneration of RuBP. The model simulation showed that photosynthetic rate and its optimal temperature were highest in summer.     

Effects of winter buds on winter nitrogen uptake and allocation in Pinus densiflora saplings

Studies of nitrogen (N) use by plants have confirmed some winter N uptake; however, the mode of regulation of plant N use in winter is unknown. The regulation of N use by plants during winter may differ from that in the growing season, as plant growth strongly affects N use. We investigated the effects of winter buds on winter N use by Japanese red pine (Pinus densiflora), as a previous study demonstrated that N absorbed during winter contributes significantly to leaf growth in the following spring. We conducted a bud pruning experiment during winter to examine the effects of winter buds on winter N uptake and allocation among plant organs using ¹⁵N labeling. Over a three-week labeling period, the ¹⁵N content in roots increased to 0.20 ± 0.12 mg N g DW^?1, which is equivalent to 1.8 ± 1.1 % of the total N content in the roots. However, this absorbed ¹⁵N rarely appeared in needles and buds. Bud pruning did not affect ¹⁵N uptake and allocation. On the other hand, significant total N retranslocation was found within the crowns of saplings without bud pruning, but N was not retranslocated in bud-pruned plants. The bud pruning experiment indicated that N was retranslocated from needles into winter buds. Since soil N availability changes dramatically and is unstable in many forest ecosystems, N contained in needles would be a more stable source of N than newly absorbed N.     

Responses of Acer saccharum canopy trees and saplings to P, K and lime additions under high N deposition

Heavy atmospheric nitrogen (N) deposition has been associated with altered nutrient cycling, and even N saturation, in forest ecosystems previously thought to be N-limited. This observation has prompted application to such forests of non-N mineral nutrients as a mitigation measure. We examined leaf gas-exchange, leaf chemistry and leaf and shoot morphological responses of Acer saccharum Marsh. saplings and mature trees to experimental additions of non-nitrogenous mineral nutrients (dolomitic lime, phosphorus + potassium (P + K) and lime plus P + K) over 2 years in the Haliburton region of central Ontario, which receives some of the largest annual N inputs in North America. Nutrients were adsorbed in the mineral soil and taken up by A. saccharum trees within 1 year of fertilizer application; however, contrary to expectation, liming had no effect on soil P availability. Saplings and canopy trees showed significant responses to both P + K fertilization and liming, including increased foliar nutrient concentration, leaf size and shoot extension growth; however, no treatment effects on leaf gas-exchange parameters were detected. Increases in shoot extension preceded increases in diameter growth in saplings and canopy trees. Vector analysis of shoot extension growth and nutrient content was consistent with sufficiency of N but marked limitation of P, with co-limitation by calcium (Ca) in saplings and by Ca, Mg and K in canopy trees.