Comparative hydraulic and anatomic properties in palm trees (Washingtonia robusta) of varying heights: implications for hydraulic limitation to increased height growth

As trees grow taller, the energetic cost of moving water to the leaves becomes higher and could begin to limit carbon gain and subsequent growth. The hydraulic limitation hypothesis states that as trees grow taller, the path length and therefore frictional resistance of water flow increases, leading to stomatal closure, reduced photosynthesis and decreased height growth in tall trees. Although this hypothesis is supported by the physical laws governing water movement in trees, its validation has been complicated by the complex structure of most tree species. Therefore, this study tested the hydraulic limitation hypothesis in Washingtonia robusta (H. Wendl.), a palm that, while growing to tall heights, is still structurally simple enough to act as a model organism for testing. There were no discernable relationships between tree height and stomatal conductance, stomatal densities, guard cell lengths, leaf dry mass per unit area (LMA) or sap flux, suggesting that these key aspects of hydraulic limitation are not reduced in taller palms. Taller palms did, however, have higher maximum daily photosynthetic assimilation rates, lower minimum leaf water potentials that occurred earlier in the day and fewer, smaller leaves than did shorter palms. Leaf epidermal cells were also smaller in taller palms compared with shorter ones. These findings are consistent with hydraulic compensation in that tall palms may be overcoming the increased path length resistance through smaller, more efficient leaves and lower leaf water potentials than shorter palms.     

Hydraulic properties of fronds from palms of varying height and habitat

Because palms grow in highly varying climates and reach considerable heights, they present a unique opportunity to evaluate how environment and plant size impact hydraulic function. We studied hydraulic properties of petioles from palms of varying height from three species: Iriartea deltoidea, a tropical rainforest species; Mauritia flexuosa, a tropical rainforest, swamp species; and Washingtonia robusta, a subtropical species. We measured leaf areas, petiole cross-sectional areas, specific conductivity (K _S), petiole anatomical properties, vulnerability to embolism and leaf water potentials and calculated petiole Huber values and leaf-specific conductivities (K _L). Leaf and petiole cross-sectional areas varied widely with height. However, hydraulic properties including Huber values, K _S and K _L, remained constant. The two palmate species, M. flexuosa and W. robusta, had larger Huber values than I. deltoidea, a pinnately-compound species which exhibited the highest K _S. Metaxylem vessel diameters and vascular bundle densities varied with height in opposing patterns to maintain petiole conductivities. I. deltoidea and W. robusta petioles had similar P ₅₀ values (the point at which 50% of hydraulic conductivity is lost) averaged over all crown heights, but W. robusta exhibited more negative P ₅₀ values in taller palms. Comparison of P ₅₀ values with transpiring midday leaf water potentials, as well as a double-dye staining experiment in a 1-m-tall palm, suggested that a fairly significant amount of embolisms were occurring and refilled on a diurnal basis. Therefore, across palms differing widely in height and growing environments, we found convergence in water transport per unit leaf area (K _L) with individuals exhibiting differing strategies for achieving this.     

Stilt Root Structure in the Neotropical Palms Iriartea deltoidea and Socratea exorrhiza1

Many arborescent palms develop a stilt root cone that provides increased mechanical support on steep hillsides, better root aeration under waterlogged conditions, resprouting after mechanical damage, and rapid stem elongation. However, for most species the role of stilt roots is not well understood. We determined how palm size and slope inclination affected the structure of stilt roots in the neotropical palms Iriartea deltoidea and Socratea exorrhiza. We expected palm height to be lower on steep slopes because the effectiveness of root support could decrease as slope inclination increases, and thus, the structure of the root cone would vary mostly with slope. Alternatively, if stilt root development is determined by palm size, their production should match stem height, with taller palms having larger cones. We measured the stilt root cone of 31 Iriartea and 36 Socratea palms in San Ramón Biological Reserve and Golfito Wildlife Refuge, Costa Rica. We divided the cone into five variables (horizontal projections of the cone base and stilt root height up and down the slope, and width of the cone base), from which we extracted the first two principal components and used them to measure the effects of slope and palm size on stilt root structure. We found stilt roots to be determined by palm size, not by slope conditions. Stilt roots matched palm size, with larger palms having taller and larger cones, maintaining stability under diverse slope conditions.     

Allometric estimation of total leaf area in the neotropical palm Euterpe oleracea at La Selva, Costa Rica

We estimated the magnitude of the total leaf area of the neotropical palm Euterpe oleracea and examined its allometry relative to the variation in stem height and diameter at La Selva Biological Station in Costa Rica. The allometric relationships between frond leaf area and frond length (from tip to base), and between frond leaf area and number of leaflets, were determined by natural logarithmic regressions to estimate the total area of each frond. Palm total leaf area was then estimated by adding the area of the composing fronds. We fit 14 separate regression models that related one or more of the morphological variables (number of fronds, diameter at breast height, stem height) to the total leaf area. Our results show that palm total leaf area is directly proportional to the number of fronds and palm size as reflected in stem height and diameter. Eight out of the 14 models had r ² values of >0.90 and incorporated a diverse combination of predictor variables. Simple linear regression models were more congruent with the observed values of total leaf area, whereas natural logarithmic models overestimated the value of total leaf area for large palms. Both approaches show a high degree of association among morphological characters in E. oleracea supporting the hypothesis that palms behave like unitary organisms, and are morphologically constrained by the lack of secondary meristems. To afford attaining canopy heights, woody palms need to show a high degree of phenotypic integration, shaping their growth and allometric relationships to match spatial and temporal changes in resources.     

BIOMASS ALLOCATION AND GEOMETRY OF THE CLONAL FOREST HERB LAPORTEA CANADENSIS: ADAPTIVE RESPONSES TO THE ENVIRONMENT OR ALLOMETRIC CONSTRAINTS?

Using a conceptual model, I predicted the direction of biomass allocation and geometric responses to several environmental variables for Laportea canadensis, a clonal forb dominating the herbaceous stratum of many North American floodplain and mesic forests. Laportea stems and plants, especially dominant ones, generally (60%) respond as predicted to canopy opening, conspecific leaf area and density, and poor drainage, but are merely reduced in growth when growing on sandier soils. However, allometric relationships explain most of the variation in geometry and allocation. Still, variation in geometry and allocation (as great as among 21 species of herbs studied by Givnish [1982]), helps explain the success of Laportea in a range of microenvironments. In upland forests, stems in canopy gaps are tallest but allocate relatively less biomass to leaves than shaded stems, suggesting that interherb competition is the major problem faced under canopy gaps. Leaf morphology also changes with increasing canopy opening—individual leaves are larger, heavier, and thicker and are displayed on more steeply ascending petioles. Floodplain plants respond to light gaps mainly with changes in leaf morphology and display. With increasing conspecific density and leaf production, Laportea stems in both uplands and floodplains grow taller, allocate relatively more biomass to stems, and display leaves higher on the stem. The allocation and geometry of taller stems are more independent of density, and more closely affected by tree-canopy opening, than are small stems. Intermediate soil textures in floodplains promote maximum Laportea production; variations in other factors are less important. Poorly drained soils in floodplains (heavy-textured soils at low elevations) cause decreased Laportea height and absolute leaf weight, but increase relative allocation to leaves and roots, as predicted. On the other hand, Laportea appears poorly adapted to sandier soils. Rather than responding to sandier soils as predicted, Laportea's overall growth is reduced. Geometric responses of Laportea to environment are mediated by allometric realities: an increase in height favored in productive environments produces a concomitant decrease in relative leaf allocation. Although predicted (presumably adaptive) shifts are significant when plant size is accounted for, most of the variation in allocation and geometry is due to allometry.     

Wet- vs. Dry-Season Transpiration in an Amazonian Rain Forest Palm Iriartea deltoidea

The wet and dry seasons in tropical rain forests can differ in precipitation, soil moisture and irradiance more significantly than often assumed. This could potentially affect the water relations of many tree species that may exhibit either increased transpiration in the dry season as a response to the increased irradiance or decreased transpiration as a result of decreases in soil moisture and increases in atmospheric vapor pressure deficit (VPD). Atmospheric data, soil moisture data and sap fluxes in Iriartea deltoidea palms were measured in eastern Ecuador during the wet and dry seasons. There were no differences between total daily sap fluxes in I. deltoidea palms during the wet and dry seasons; however, evaporative demand was significantly higher in the dry season and therefore, transpiration was more restricted by stomatal closure during the dry season than the wet season. This is likely the result of larger atmospheric VPD during the dry season compared with the wet season and possibly the result of reduced soil moisture availability. Additionally, based on published tree abundances in this area, measured sap fluxes in I. deltoidea were scaled up to the hectare level. Transpiration from I. deltoidea palms was estimated to be around 0.03 mm/d, which could represent about 1 percent of total transpiration in this area of the Amazon rain forest. If climate change predictions for more lengthy tropical dry periods are realized, greater stomatal control of dry-season sap flux has the potential to become even more prevalent in tropical species.     

Changes in leaf morphology and anatomy with tree age and height in the broadleaved evergreen species, Eucalyptus regnans F. Muell

Relatively little is known about changes in leaf attributes over the lifespan of woody plants. Knowledge of such changes may be useful in interpreting physiological changes with age. This study investigated changes in leaf morphology and anatomy with tree age and height in the broadleaved evergreen species, Eucalyptus regnans. Fully expanded leaves were sampled from the upper canopy of tree ages ranging from 6 to 240 years, and tree heights ranging from about 10–80 m. There were significant changes in leaf form with increasing tree age and height. Leaf size and specific leaf area (SLA; leaf area/leaf mass) decreased, leaf thickness increased, and leaves became narrower relative to their length, with increasing tree age and height. Cuticle thickness and leaf waxiness, including wax occlusion of the stomatal antechamber, increased with increasing age and height. By comparison, there were no clear trends in stomatal frequency or stomatal length with tree age, although there were curvilinear relationships between an index of total stomatal pore area per leaf lamina and both tree age and tree height. The results support the hypothesis that leaves of E. regnans become more xeromorphic with tree age and height. The results are discussed in relation to their significance for changes in water relations in the canopy with age.     

Population Structure and Life Cycle of Borassus aethiopum Mart.: Evidence of Early Senescence in a Palm Tree1

As with other plants having a relatively simple morphology, solitary palms are useful biological models for studying the life histories of long-lived plants. In the first study to investigate the life history of Borassus aethiopum, a widespread dioecious palm growing in African savannas, we found that: (1) the number of leaves increased up to reproductive maturity and then decreased during the reproductive period, while height increased throughout life; (2) female fecundity, measured as the number of seedlings within a few meters under the female canopies, decreased markedly in old individuals with few leaves; and (3) height distribution was strongly bimodal. This bimodality was due to variations in the stem elongation rate during the life of the palm and the accumulation of adults with low mortality rates in the taller height classes. The observed pattern of fecundity and number of leaves in relation to height clearly suggests a senescence period that began just after sexual maturity and appeared to be protracted. Comparisons between the life history of B. aethiopum and the life histories of some forest palms showed that environmental conditions cannot in themselves explain the various palm life histories.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系,其异速生长指数<1.0,但该假说是否适用于不同生活型（常绿与落叶）亚热带木本植物不同冠层高度（上下冠层）当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象,采用标准化主轴回归估计（standardized major axis estimation,SMA）方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示：（1）当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异（P>0.05）,叶面积在常绿和落叶植物中有显著差异（P<0.05）,常绿和落叶植物的比叶重存在显著差异（P<0.05）,而落叶植物的比叶重在不同冠层高度存在显著差异（P<0.05）,同一冠层,常绿植物比叶重显著高于落叶植物（P<0.05）;（2）69种植物的叶面积与叶生物量异速生长指数具有物种特异性,60.9%的物种叶面积与叶生物量呈等速生长关系;（3）不同冠层和生活型植物的叶面积与叶生物量呈等速生长关系,但其异速生长常数在不同冠层高度与生活型间存在差异。这些结果表明冠层高度和生活型未改变叶面积-生物量之间的等速生长关系,不支持"收益递减"假说。     

Leaf structure and patterns of photoinhibition in two neotropical palms in clearings and forest understory during the dry season

We studied the leaf structural, water status, and fast fluorescence responses of two palms, Socratea exorrhiza and Scheelea zonensis, under natural dry season conditions in a clearing (high light [HL] palms) and the forest understory (low light [LL] palms) on Barro Colorado Island, Panama. HL-Socratea leaves were more shade-adapted, less xeromorphic, and more strongly affected by drought than HL-Scheelea. F_v/F_m (the ratio of variable to maximum chlorophyll fluorescence) and t_½ (the half-rise time of F_m) was lower in HL-leaves of both species, indicating photoinhibition. In HL-Scheelea, the light-induced reduction of F_v/F_m was much less than in HL-Socratea, and F_v/F_m recovered completely overnight. Patterns of relative water content, specific leaf dry weight, stable carbon isotope composition, and leaf conductance suggest that increased drought resistance in Scheelea reduces susceptibility to photoinhibition. An increase in F_o indicated the inactivation of PSII reaction centers in HL-Socratea. The very low chlorophyll a/b ratio and alterations in chloroplast ultrastructure in HL-Socratea are consistent with photoinhibition. Under LL, the species showed no appreciable interspecific differences in chlorophyll fluorescence. Excess light leads to low values of F_v/F_m in HL-plants relative to LL-plants on both leaf surfaces, particularly on the lower surface, due to a decrease of F_m in both surfaces and an increase in F., of lower surface. For both species, F_o for the lower surfaces of HL-plants was higher and t_½ was markedly lower than for the upper surface, as is typical for shade-adapted leaves. Xeromorphic leaf structure may reduce susceptibility to photoinhibition during the dry season. Drought-enhanced photoinhibition could limit the ability of some species to exploit treefall gaps.     

Root (wilt) disease of coconut palms in South Asia – an overview

Root (wilt) disease (RWD) caused by phytoplasma is one of the most devasting diseases of coconut palms. The major symptoms of the disease in leaves are wilting and drooping and flaccidity; ribbing, paling/yellowing and necrosis of leaflets are typical symptoms of foliar diseases. Unopened pale yellow leaflets of spindle leaves are more susceptible to leaf rot disease, which is caused by Exerohilum rostratum and Colletotrichum gloeosporioides. RWD is caused by phytoplasmas, the cell wall-less prokaryotes that are bounded by a “unit” membrane. In ultrathin sections, they appear as a complex multi-branched, beaded, filamentous or spheroidal pleomorphic bodies. The disease was transmitted by plant hoppers (Proutista moesta) and lace wing bug (Stephanitis typica). Phytoplasmas are generally present in the phloem sieve tubes and in the salivary glands of these insect vectors. Phytoplasmas cannot be cultured in vitro, and hence it is very difficult to identify them. Using polymerase chain reaction technique, group-specific primers have been applied to detect mixed-phytoplasma infections in a single host. RWD, is a non-lethal, debilitating disease, and hence an integrated approach for the management of this disease in coconut palms has been discussed in this study.     

Characterization of X-ray induced increase of mitotic cross-overs in Glycine max

Summary Three types of spots can be identified on the leaves of heterozygous light green, Y/y, Glycine max (L.) Merrill: dark green (D) and aurea (A) single spots (resembling the phenotypes of the homozygotes) and double (Db) spots consisting of adjacent D and A tissue. X-irradiation increased the frequency of each type of spot on simple and first compound leaves. The Db spots, indicative of mitotic crossing-over (MCO), increase linearly with increasing dosage. Moisture content of the seeds was independent of the rate of spot increase. At high dosages morphological alterations were observed, including spots on homozygotes, leaf area reduction, smaller seedlings, and abnormal leaf shapes. The frequency of light green spots on normal dark green, Y/Y, seedlings was tabulated and, as with all other spot types, increased with increasing X-ray dosage. Dormant soybean seeds contain leaf primordia of both simple and first compound leaves. Mature simple leaves contained more spots, reflecting a larger primordial cell number, while first compound leaves had larger spots, since each affected cell underwent more mitoses prior to leaf maturation. Within first compound leaves, the terminal leaflets developed asynchronously in relation to the lateral leaflets. Terminal leaflets were shown to be initiated first, have a larger percentage of the leaflet area covered with spots, and have larger mature leaflet area. The spontaneous rate of MCO, 3.39×10^–5 MCO events per mitosis, was increased 282-fold by 1600 R. We also ascertained that Mitomycin C is more specific for Db spot induction than X-rays. These results are compared with our similar irradiation experiments on tobacco shoot apices.     

Monocot leaves are eaten less than dicot leaves in tropical lowland rain forests: correlations with toughness and leaf presentation

Background and Aims

In tropical lowland rain forest (TLRF) the leaves of most monocots differ from those of most dicots in two ways that may reduce attack by herbivores. Firstly, they are tougher. Secondly, the immature leaves are tightly folded or rolled until 50–100 % of their final length. It was hypothesized that (a) losses of leaf area to herbivorous invertebrates are generally greatest during leaf expansion and smaller for monocots than for dicots, and (b) where losses after expansion are appreciable any difference between monocots and dicots then is smaller than that found during expansion.

Methods

At six sites on four continents, estimates were made of lamina area loss from the four most recently mature leaves of focal monocots and of the nearest dicot shoot. Measurements of leaf mass per unit area, and the concentrations of water and nitrogen were made for many of the species. In Panama, the losses from monocots (palms) and dicots were also measured after placing fully expanded palm leaflets and whole dicot leaves on trails of leaf-cutter ants.

Key Results

At five of six sites monocots experienced significantly smaller leaf area loss than dicots. The results were not explicable in terms of leaf mass per unit area, or concentrations of water or nitrogen. At only one site was the increase in loss from first to fourth mature leaf significant (also large and the same in monocots and dicots), but the losses sustained during expansion were much smaller in the monocots. In the leaf-cutter ant experiment, losses were much smaller for palms than for dicots.

Conclusions

The relationship between toughness and herbivory is complex; despite the negative findings of some recent authors for dicots we hypothesize that either greater toughness or late folding can protect monocot leaves against herbivorous insects in tropical lowland rain forest, and that the relative importance varies widely with species. The difficulties of establishing unequivocally the roles of leaf toughness and leaf folding or rolling in a given case are discussed.Key words: anti-herbivore defences, dicots, herbivory, leaf folding, leaf rolling, leaf toughness, monocots, palms, tropical rain forest     

Effect of Irradiance, Sugars and Nitrogen on Leaf Size of In Vitro Grown Ceratonia Siliqua L.

Carob (Ceratonia siliqua L.) has compound pinnate leaves consisting of 4 – 6 pairs of leaflets. However, in conditions of in vitro culture only one pair of leaflets develops. With increasing irradiance from 9.3 to 74.1 µmol m^–2 s^–1, leaf area increased 5-fold. Sucrose also significantly increased leaf area and the maxima were at concentration 147 mM at high irradiance and 233.6 mM at low irradiance. Sucrose was superior to fructose, glucose and combination of both in increasing leaf area. Decreasing concentration of KNO₃ and NH₄NO₃ caused a 3-fold decline of leaf area.     

Morphological,structural and physiological differences in heteromorphic leaves of Euphrates poplar during development stages and at crown scales

• Euphrates poplar (Populus euphratica Oliv.) has heteromorphic leaves including strip, lanceolate, ovate, and broad‐ovate leaves from base to top in the mature canopy.
• To clarify how diameter at breast height (DBH) and tree height affect the functional characteristics of all kinds of heteromorphic leaves, we measured the morphological anatomical structure and physiological indices of five crown heteromorphic leaves of P. euphratica at 2, 4, 6, 8, 10, and 12 m from the same site. We also analysed the relationships between morphological structures and physiological characteristics of heteromorphic leaves and DBH and the height of heteromorphic leaves.
• The results showed that the number of abnormalities regarding blade width, leaf area, leaf thickness, leaf mass per area, cuticle layer thickness, palisade tissue thickness, and palisade tissue/sponge tissue ratio increased with size order and sampling height gradient. Net photosynthetic rate, transpiration rate, stomatal conductance, instantaneous water use efficiency, stable delta carbon isotope ratio, proline and malondialdehyde (MDA) content increased with DBH and sampling height. By contrast, blade length, leaf shape index, and intercellular CO2 concentration decreased with the increase in path order and sampling height gradient. Although MDA content and leaf sponge thickness were not correlated with DBH or sampling height, other morphological structure and physiological parameters were significantly correlated with these variables. In addition, correlations were found among leaf morphology, anatomical structure, and physiological index parameters indicating that they changed with path order and tree height gradient.
• The differences in the morphology, anatomic structure and physiological characteristics of the heteromorphic leaves ofP. euphratica are related to ontogenesis stage and coronal position.

     

Light partitioning among species and species replacement in early successional grasslands

Abstract. We studied canopy structure, shoot architecture and light harvesting efficiencies of the species (photon flux captured per unit above‐ground plant mass) in a series of exclosures of different age (up to 4.5 yr) in originally heavily grazed grassland in N Japan.Vegetation height and Leaf Area Index (LAI) increased in the series and Zoysia japonica, the dominant in the beginning, was replaced by the much taller Miscanthus sinensis. We showed how this displacement in dominance can be explained by inherent constraints on the above‐ground architecture of these two species. In all stands light capture of plants increased with their above‐ground biomass but taller species were not necessarily more efficient in light harvesting. Some subordinate species grew disproportionally large leaf areas and persisted in the shady undergrowth. Some other species first grew taller and managed to stay in the better‐lit parts of the canopy, but ultimately failed to match the height growth of their neighbours in this early successional series. Their light harvesting efficiencies declined and this probably led to their exclusion. By contrast, species that maintained their position high in the canopy managed to persist in the vegetation despite their relatively low light harvesting efficiencies. In the tallest stands ‘later successional’ species had higher light harvesting efficiencies for the same plant height than ‘early successional’ species which was mostly the result of the greater area to mass ratio (specific leaf area, SLA) of their leaves. This shows how plant stature, plasticity in above‐ground biomass partitioning, and architectural constraints determine the ability of plants to efficiently capture light, which helps to explain species replacement in this early successional series.     

Shell Morphological Variation of Littoraria angulifera among and within Mangroves in NE Brazil

The shell morphological variation of the periwinkle Littoraria angulifera (Lamarck, 1822) was studied in tropical northeast Brazilian mangroves. This area was selected because mangroves in different stages of regeneration, and thus different tree heights can be found. We evaluated whether differences in solar radiation due to differences in tree height influenced the distribution and shell morphology of L. angulifera, and carried out an experiment to test if individuals collected from mangroves with different tree heights differed in their resistance to desiccation. We also analysed if there were differences in L. angulifera shell length and shape between tidal levels within a mangrove. Finally, we tested if increased habitat complexity due to the presence of oysters could influence L. angulifera shell length and shape in different-statured mangroves. We predicted that the oysters could reduce desiccation stress on periwinkles in small-statured mangroves but have no effect in taller ones. Shell length and shape varied among mangroves, although a large variation within mangroves was also recorded. Shell proportionality (shell length:width ratio) increased with shell length, and this relationship differed among mangroves; however, no differences were found in a subsequent year. Individuals from small-statured mangroves survived longer than those from taller mangroves in the desiccation experiment, with a weak correlation between shell proportionality and loss of mass. The presence of oysters had no apparent influence on shell morphology that could be correlated with reduced desiccation stress. The patterns found varied greatly both at small and large spatial scales, suggesting that future studies should evaluate phenotypic and genetic variation at the same time to properly understand variation in L. angulifera shell morphology.     

平茬高度对香椿生长及其光合特性和非结构性碳水化合物含量的影响

该研究以4年生香椿为试验材料,设置平茬20 cm(T₁)、50 cm(T₂)、80 cm(T₃)和不平茬(CK)4种处理,观测其萌枝和叶片生长情况,以及叶片的气体交换参数、光合色素含量、非结构性碳水化合物(NSC)含量的变化,分析不同平茬高度的生长生理响应差异,以明确平茬措施下香椿植株更新复壮再生的生理机制。结果表明：(1)平茬能够显著提高香椿的萌枝能力,促进侧枝和叶片生长,其萌枝数、侧枝长度在T₃处理下最高,成枝数、叶长、叶宽、叶面积及侧枝粗度在T₂处理下达到最大值。(2)随着平茬高度的增加,香椿叶片净光合速率、蒸腾速率、气孔导度和水分利用效率均先升后降,并在T₂处理达到最大,较CK分别显著提高了17.33%、10.00%、13.51%和6.98%;平茬也提高了叶片光合色素含量,叶绿素a、叶绿素b、总叶绿素含量和类胡萝卜素含量在T₂处理下分别比CK显著提高了18.34%、27.07%、21.11%和23.05%。(3)不同平茬高度处...     

Changes in foliage distribution with relative irradiance and tree size: Differences between the saplings ofAcer platanoides andQuercus robur

Dependencies of foliage arrangement and structure on relative irradiance and total height (TH) were studied in saplings ofAcer platanoides andQuercus robur. The distribution of relative foliar area and dry weight (leaf area and weight in a crown layer per total tree leaf area and weight, respectively) were examined with respect to relative height (RH, height in the crown per TH) and characterized by the Weibull function. The distributions of relative area and weight were nearly identical, and the differences between them were attributable to a systematic decline in leaf dry weight per area with increasing crown depth. Foliage distribution was similarly altered by tree size in both species; RH at foliage maximum was lower and relative canopy size (RCS, length of live crown per TH) greater in taller trees. However, the distribution was more uniform inA. platanoides than inQ. robur. Apart from the size effects, relative irradiance also influenced canopy structure; RCS increased inQ. platanoides and decreased inQ. robur with increasing irradiance. As crown architecture was modified by irradiance, foliage distribution was shifted upwards with decreasing irradiance inA. platanoides, but it was independent of irradiance inQ. robur. Higher foliage maximum at lower irradiance in more shade-tolerantA. platanoides is likely to contribute towards more efficient foliar display for light interception and increase the competitive ability of this species in light-limited environments. Consequently, these differences in crown architecture and foliage distribution may partly explain the superior behavior ofA. platanoides in understory.     

长白山红松不同树高处径向生长特征及其对气候的响应

利用长白山红松不同树高(0.3、1.3、4、10、15、20、25 m)处的径向生长资料,分析各树高处径向生长特征,建立红松生长与气候因子的相关关系,以期完善红松种群对气候变化的响应机制。结果表明:(1)红松不同树高处年径向生长量变化趋势基本一致,除在1980年前后,20 m处径向生长量出现异常增加外,其他各高度径向生长均出现下降趋势,红松基部和顶端(0.3、1.3 m和20 m)处径向生长年际变化更明显。随着树高增加,各处年径向生长率有所降低,0.3m处生长速率最大,且与10 m和15 m处径向生长差异显著(P < 0.05)。(2)不同树高处径向生长对气候因子的响应存在明显差异,10 m树高是红松径向生长对温度和降水响应差异的分界线。10 m以下红松径向生长主要受到生长季温度的负作用,尤其是4 m处,与当年生长季初期(4月和5月)温度显著负相关(P < 0.05)。0.3 m和1.3 m处径向生长分别与上年9月平均温度显著正相关(P < 0.05),当年6月平均和最高温度显著负相关(P < 0.05)。随着树高上升,降水对径向生长的促进作用增强,而温度对径向生长的作用也发生改变。10 m(含)以上则受到温度和降水的共同作用。10 m处径向生长对气候因子响应最敏感,受到当年生长季高温的抑制作用,还与上年和当年生长季末(9月)降水显著正相关(P < 0.05)。15 m处径向生长与上年9月最低温度和降水显著正相关(P < 0.05),而与当年5月月平均温度显著负相关(P < 0.05)。20 m处径向生长与当年3月月平均、最低和最高温度,当年7月月平均温度以及当年5月降水显著正相关(P < 0.05),而与当年1月降水显著负相关(P < 0.05)。