Quantitative analysis of polymeric proanthocyanidins in birch leaves with normal-phase HPLC

The proanthocyanidin composition and content in the leaves of nine birch species (Betula albosinensis, B. ermanii B. maximowicziana, B. nana, B. papyrifera, B. pendula, B. platyphylla, B. pubescens, and B. pubescens ssp. czerepanovii) were studied with different methods including colorimetric assay, HPLC coupled with PAD or ESI/MS and NMR. Total proanthocyanidin content was determined using the acid butanol assay. A normal phase-HPLC method was applied for the analysis of polymeric proanthocyanidins. The content of polymeric proanthocyanidins was estimated from a late eluting peak in the chromatogram. With this HPLC method, quantitative analysis of polymeric proanthocyanidins could be performed directly from leaf extracts: no additional purification or preparation steps were required. It was shown that birch leaves contained mainly polymeric proanthocyanidins with a degree of polymerisation greater than 10. Total proanthocyanidin content (expressed as dry weight) was found to vary from 44mg/g (B. papyrifera) to 145mg/g (B. nana), and polymeric proanthocyanidin content from 39mg/g (B. pendula) to 119 mg/g (B. nana).     

不同海拔长白山岳桦的生理变化

通过分析长白山国家级自然保护区内不同海拔(A1:1700 m, A2:1800 m, A3:1900 m, A4:2000 m, A5:2050 m)梯度岳桦叶片中各种生理指标含量的变化,探讨了林线树木适应高山环境的生理机制.结果表明:随着海拔的升高,比叶面积(SLA)显著减小,A5与A1相比下降了35.90%,差异达到显著水平;叶绿素含量随海拔梯度升高而降低,但叶绿素a/b比值(Chla/Chlb)和Car的相对含量(Car/Chl)随海拔梯度升高而增加;在海拔1900 m左右,MDA含量和MP均处于最低水平,各种酶的活性均为最低;当海拔超过2000 m,接近森林分布的界限时,MDA含量和MP升高,并达到最大值,各种酶的活性都出现了一定程度的下降.综合本次研究表明, 在海拔1900 m比较适合岳桦的生长;海拔超过2000 m,岳桦体内生理抗性下降,不利于岳桦的生长发育,因此高海拔限制了岳桦的分布.     

Control of Leaf Growth and its Role in Determining Variation in Plant Growth Rate from an Ecological Perspective

Abstract: Plants vary widely in their relative growth rate (RGR), be it dependent on environmental conditions or due to their genetic background. In a comparison of the RGR of grasses growing under different environmental conditions, variation in RGR tends to correlate with that in the leaf elongation rate (LER). When different species or genotypes thereof are compared under identical growing conditions, variation in LER may or may not correlate with that in RGR, depending on the comparison. However, since RGR is described by an exponential equation, whereas LER is mainly a linear process, we conclude that any correlation between RGR and LER must be fortuitous. That is, exponential growth must be due to increases with time in plant traits such as 1) leaf dry mass per unit leaf length invested per unit time, and/or 2), i.e., the total LER of all the growing leaves at one point in time. The latter can be achieved as follows: 1) each subsequent leaf has a higher LER than the preceding one; 2) leaves appear at an increasing rate; 3) the duration of the process of leaf elongation increases for subsequent leaves. In this review, we only explore possible factors that account for changes in with time, in different genotypes and under different environmental conditions. Inherent variation in LER of individual leaves and variation due to environmental factors may reflect variation in the rate of cell division and/or in cell elongation.     

Effects of Manganese in Solution Culture on the Growth of Five Deciduous Broad-Leaved Tree Species with Different Successional Characters from Northern Japan

The effects of four manganese (Mn) concentrations (1, 10, 50, and 100 g m^-3 = Mn₁, Mn₁₀, Mn₅₀, Mn₁₀₀) in solution culture on growth variables were studied for seedlings of five deciduous broad-leaved trees with different successional characteristics and shoot development patterns in northern Japan. The five species were: Betula ermanii, Betula platyphylla var. japonica, and Alnus hirsuta (early-successional species with continuous leaf development), Ulmus davidiana var. japonica (mid-successional species with flush and continuous leaf development), and Acer mono (late-successional species with a flush type leaf development). In plants grown in the Mn environment for about 45 d, relative growth rate (RGR) decreased with increasing Mn supply. Between the 1 and 100 g(Mn) m^-3, RGR decreased by 20 % for B. ermanii and B. platyphylla, by 40 % for A. hirsuta and A. mono, and by 80 % for U. davidiana. Specific leaf area (SLA) and leaf mass ratio (LMR) of all species were little affected by high Mn supply. In U. davidiana, however, there was a 67 % decrease in LMR in Mn₁₀₀ plants. Leaf area ratio (LAR) was higher in early-successional species than in mid- and late-successional ones but differed little among Mn treatments within species, except for U. davidiana where LAR declined substantially with increased Mn supply. While LAR, which represents the relative size of assimilatory apparatus, was little affected, net photosynthetic rate (P_N) saturated with radiant energy decreased with increasing Mn supply in all species. Thus P_N was adversely affected by high accumulation of Mn in leaves, which resulted in an overall reduction in biomass production. However, the proportional allocation of photosynthates to the assimilatory apparatus was not affected by different Mn toxicity in hardwood tree seedlings. This revised version was published online in September 2006 with corrections to the Cover Date.     

Shoot morphogenesis associated with flowering in Populus deltoides (Salicaceae)

Temporal and spatial formation and differentiation of axillary buds in developing shoots of mature eastern cottonwood (Populus deltoides) were investigated. Shoots sequentially initiate early vegetative, floral, and late vegetative buds. Associated with these buds is the formation of three distinct leaf types. In May of the first growing season, the first type begins forming in terminal buds and overwinters as relatively developed foliar structures. These leaves bear early vegetative buds in their axils. The second type forms late in the first growing season in terminal buds. These leaves form floral buds in their axils the second growing season. The floral bud meristems initiate scale leaves in April and begin forming floral meristems in the axils of the bracts in May. The floral meristems subsequently form floral organs by the end of the second growing season. The floral buds overwinter with floral organs, and anthesis occurs in the third growing season. The third type of leaf forms and develops entirely outside the terminal buds in the second growing season. These leaves bear the late vegetative buds in their axils. On the basis of these and other supporting data, we hypothesize a 3-yr flowering cycle as opposed to the traditional 2-yr cycle in eastern cottonwood.     

Geographical variation and population structure in the White-rumped Sandpiper Calidris fuscicollis as shown by morphology, mitochondrial DNA and carbon isotope ratios

Several plant-herbivore hypotheses are based on the assumption that plants cannot simultaneously allocate resources to growth and defence. We studied seasonal patterns in allocation to growth and putatively defensive compounds by monitoring several chemical and physical traits in the leaves of mountain birch from early June (budburst) to late September (leaf senescence). We found significant seasonal changes in all measured characteristics, both in terms of concentrations (mg g^-1) and amounts (mg leaf^-1). Changes were very rapid in the spring, slow in the middle of the season, and there was another period of fast changes in the senescing leaves. Co-occurring changes in physical leaf traits and concentrations of several compounds indicated a seasonal decline in foliage suitability for herbivores. Concentrations of protein and free amino acids declined through the growing season whereas individual sugars showed variable seasonal patterns. The seasonal trends of phenolic groups differed drastically: concentrations of soluble proanthocyanidins increased through the season, whereas cell wall-bound proanthocyanidins, gallotannins and flavonoid glycosides declined after an initial increase in young leaves. We failed to find proof that the seasonal accumulation of phenolics would have been seriously compromised by leaf or shoot growth, as assumed by the growth/differentiation balance hypothesis and the protein competition model hypothesis. On the contrary, there was a steady increase in the total amount of phenolics per leaf even during the most active leaf growth.     

Resistance to the birch leafminer Fenusa pusilla (Hymenoptera: Tenthredinidae) within the genus Betula

Thirteen Betula species were tested for resistance to the birch leafminer, Femusa pusilla (Lepeletier), using no-choice assays. Birch leafminers were able to oviposit into expanding leaves of all Betula individuals tested. Larvae did not survive within any of the tested individuals of three species, B. alleghaniensis (Britt.), B. grossa (S. & Z.), and B. lenta (L.). Leafminer eggs deposited into the leaves of these species hatched, and larvae fed for a short period before dying. These three species were classified as highly resistant to birch leafminer, based on very low percent of mines (0.6-3.1%) with a diameter >3 mm. Eight species, B. papyrifera (Marsh), B. pendula (Roth), B. turkestanica (Litvin), B. glandulifira (Regal), B. ermanii (Cham.), B. platyphylla variety japonica [(Miq.) Hara], B. populifolia (Marsh) and B. maximowicziana (Regal) were classified as susceptible, with percent of mines >3 mm diameter of 87-94%. Two species, B. costata (Trautv.) and B. davurica (Pall.), displayed intermediate and variable resistance. B. davurica exhibited a mechanism of resistance not observed in the other species, Eggs oviposited into the leaves of resistant B. davurica individuals became surrounded by an area of discolored and necrotic tissue, and died. This response resembles the programmed cell death associated with a hypersensitive response.     

Trade‐offs between constitutive and induced defences drive geographical and climatic clines in pine chemical defences

There is increasing evidence that geographic and climatic clines drive the patterns of plant defence allocation and defensive strategies. We quantified early growth rate and both constitutive and inducible chemical defences of 18 Pinaceae species in a common greenhouse environment and assessed their defensive allocation with respect to each species' range across climatic gradients spanning 31^o latitude and 2300 m elevation. Constitutive defences traded‐off with induced defences, and these defensive strategies were associated with growth rate such that slow‐growing species invested more in constitutive defence, whereas fast‐growing species invested more in inducible defence. The position of each pine species along this trade‐off axis was in turn associated with geography; moving poleward and to higher elevations, growth rate and inducible defences decreased, while constitutive defence increased. These geographic patterns in plant defence were most strongly associated with variation in temperature. Climatic and geographical clines thus act as drivers of defence profiles by mediating the constraints imposed by trade‐offs, and this dynamic underlays global patterns of defence allocation.     

Induced resistance in mountain birch: defence against leaf-chewing insect guild and herbivore competition

Summary The effect of leaf damage simulating the feeding of early season insect herbivore species, e.g. Epirrita autumnata, to mountain birch, Betula pubescens ssp. tortuosa, on the performance of insect larvae was studied with eleven leaf-chewing sawfly species. I found variation in the results that was due to short- and long-term inducible responses and to the phenology of herbivore species. In general, early and mid-season species were more strongly affected by induced reactions than late-season species. This finding is in accordance with earlier results but I could show that the persistance of induced reactions rather than the influence of timing of damage is responsible for the result. The growth of the larvae of mid-season sawfly species was affected by both short- and long-term induced reactions. This result shows that early season species may escape short-term induced reactions of mountain birch in current year but may not avoid long-term effects. It is supposed that seasonal deterioration of leaf quality either masks the effects of induced defences or late-season species are better adapted to low-quality leaves. Some species show variation in their response to induced defence in different years. This may be due to yearly differences in induced reactions as well as to species-specific responses. Induced defence reactions may play a role in competitive interactions between herbivore species in leaf-chewing guild of mountain birch.     

Root herbivore performance suppressed when feeding on a jasmonate‐induced pasture grass

1. Plants defend themselves from insect herbivore attack using a range of physical and chemical defences which are in many cases regulated by phytohormones such as jasmonates. While much more is known about how jasmonates regulate defence against above‐ground herbivores (e.g. herbivores of leaves), there is increasing interest in how they influence below‐ground defences. 2. For the Poaceae, most below‐ground studies focus on highly domesticated cereals. Here it is demonstrated how exogenous application of methyl jasmonate (MeJA) to the leaf blades of a non‐domesticated pasture grass (Microlaena stipoides) caused a more than two‐fold decrease in relative growth rate (RGR) of a root‐feeding chafer (Dermolepida albohirtum). MeJA treatment did not affect root consumption rates, but substantially reduced the efficiency of conversion of ingested food to body mass. 3. Non‐targeted metabolomics identified significant changes in the metabolome of MeJA‐induced plants, with three compounds (a galactolipid, a trihydroxy fatty acid and a lysophospholipid) found to be correlated with herbivore RGR, although their roles in herbivore defence remain uncertain. 4. This study suggests that an important Australian pasture grass can become better defended against root herbivores via enhanced jasmonate activity.     

长白山自然保护区北坡岳桦树根系内生真菌的种类及分布

为研究长白山自然保护区北坡岳桦树（Betula ermanli）的根系内生真菌的种类及其分布,笔者于2011年7～9月分别在长白山自然保护区北坡海拔1700～2100m处采集岳桦树根系组织。分离得到411株内生真菌,经形态学鉴定,分别属于17个属。研究结果表明：除不产孢菌株外,青霉属Penieillium分离频率最高（21．90％）,木霉属Trichodmma分离频率次之（17．52％）,镰孢菌属Fusarium、伞形霉属Umbelopsis和曲霉属Aspergillus分离频率也相对较高（5．60％、5．35％和4．87％）。通过分析发现,从不同月份和不同海拔高度采集的岳桦树根系分离的内生真菌的种类和数量不同。试验结果表明,长白山自然保护区岳桦树根系的内生真菌种类较多,其种类、数量和分布与采样时间以及海拔等因素存在一定的联系。     

Maximum Fruit Growth Potential Following Resource Limitation During Peach Growth

To achieve its maximum organ growth potential, an organ mustgrow at its potential relative growth rate (RGR) throughoutdevelopment. When resource availability limits growth, the RGRis reduced below the potential RGR. This study examines whether,following a period of resource-limited growth, the RGR is ableto increase to the potential RGR when sufficient resources areavailable. Fruit RGRs of a late maturing peach cultivar wereexamined following removal of most of the fruits (heavy thinning)from previously unthinned trees in Apr., May, and Jun. The fruitRGRs after imposition of the thinning treatments were higherthan those on unthinned trees during source-limited periodsof the growing season, suggesting that fruit RGR can increasein response to increased resource availability. In general,the RGRs of fruits of trees thinned in Apr., May, and Jun. didnot exceed those of fruits on trees thinned at bloom, suggestingthat heavy thinning at bloom provides a reasonable estimateof the potential RGR. There were times, however, when the effectsof competition with vegetative sinks were apparent, suggestingthat the RGR of fruits on trees that were heavily thinned atbloom may underestimate the potential RGR during these times.The absolute growth rates of fruits on thinned trees were greaterthan those on unthinned trees, but generally were not greaterthan those on trees that were thinned at bloom, suggesting thatpeach fruits are unable to recover potential growth lost duringresource-limited growth periods.Copyright 1995, 1999 AcademicPress Prunus persica (L.) Batsch, peach, maximum fruit growth potential, relative growth rate, absolute growth rate, thinning, fruit-fruit competition, resource availability, resource limitation, growth analysis     

Comparison of leaf life span, photosynthesis and defensive traits across seven species of deciduous broad-leaf tree seedlings

BACKGROUND AND AIMS: Leaf life span, photosynthetic parameters and defensive traits were compared across seven species of deciduous broad-leaved tree seedlings native to northern Japan to test the "cost-benefit hypothesis" that more productive leaves are more susceptible to herbivore attack than less productive leaves. METHODS: Studies were made on three early successional species, Alnus hirsuta, Betula maximowicziana and Betula platyphylla "japonica"; one mid-successional species, Ostrya japonica, and three late-successional species, Carpinus cordata, Quercus mongolica 'grosseserrata' and Acer mono. Photosynthetic parameters and defensive traits (total phenolics, condensed tannin and toughness) of leaves were measured for each species, and a bioassay test with Eri silkmoth larvae (Samia cynthia ricini) was undertaken to evaluate differences between species in susceptibility to herbivore attack. KEY RESULTS: Early successional species have a shorter leaf life span (62-88 d) than late successional species (155-187 d). Leaf nitrogen content and light-saturated photosynthetic rate per unit leaf area (P(sat)-area) and per unit leaf mass (P(sat)-mass) were negatively correlated with leaf life span. The nitrogen content of early successional species was about 30 mg g(-1) and that of late successional species was about 16 mg g(-1). Leaf toughness and the C/N ratio were positively correlated with leaf life span, although condensed tannin was not correlated with leaf life span. The bioassay test showed that the number of days the larvae survived was negatively correlated with leaf life span. Average survival of larvae feeding on leaves of A. hirsuta, which has the shortest leaf life span, was 14.4 d and that of Q. mongolica, which has the longest leaf life span, was 6.6 d. The number of days of larval survival was positively correlated with leaf nitrogen content. There was no correlation between days of larval survival and defensive traits. CONCLUSIONS: These results indicate that species with a shorter leaf life span have higher photosynthetic productivity and are more susceptible to herbivore attack than species with a longer leaf life span. This supports the "cost-benefit hypothesis".     

岳桦幼苗光合特性和非结构性碳水化合物积累对干旱胁迫的响应

以长白山林线树种岳桦为对象,利用生长控制试验进行干旱处理,研究干旱对岳桦幼苗光合特性及非结构性碳水化合物(NSC)积累的影响.结果表明:干旱显著降低了岳桦幼苗的净光合速率和气孔导度,提高了其水分利用效率;干旱显著增加了岳桦幼苗叶、皮、干和根中的可溶性糖和总NSC的含量,但显著降低了淀粉含量;随着干旱的持续,叶片的气孔导...     

Growth characteristics of root-shoot relations of three birch seedlings raised under different water regimes

Three birch species (Betula ermanii, B. maximowicziana, B. platyphylla var. japonica) widespread in northern Japan were raised under different water regimes (ca. 23, 35, and 60%) to study root-shoot increment and gas exchange traits in relation to their habitat preferences in natural conditions. Total biomass of all birches was larger for medium, wet and dry treatment. Maximum root length of B. platyphylla raised at dry or wet condition was higher than other species. Root growth rate of three birches peaked around mid July to early September while shoot growth rate was found maximum between late June and mid August. Root growth of three birches was suppressed under dry and wet treatment, especially for B. ermanii. Allocation percentage of biomass to roots of three birches ranged from 30 to 40% but it mainly allocated to elongate the root length under dry and wet condition. Fine roots of B. ermanii and B. maximowicziana under wet condition were distributed mainly on soil surface. In the dry treatment, B. platyphylla allocated photosynthates to elongate the root length and fine root production (<2 mm) and had highest plasticity of roots to different water conditions among the three birches. Specific leaf area (SLA) of three birches decreased with decreasing soil moisture content. Plasticity in SLA of B. platyphylla was found largest. Net photosynthetic rate (Pn) and water use efficiency (WUE) of B. platyphyllashowed highest among all the treatments. Leaf nitrogen content of B. platyphyllawas also relatively higher under different water conditions as compared to the other two birch species, suggesting B. platyphylla may enable to invade to various growing conditions with different water regimes.     

Modelling seedling growth rates of 18 temperate arable weed species as a function of the environment and plant traits

BACKGROUND AND AIMS: The early growth rate of seedlings in the exponential phase is an important eco- physiological trait in crop/weed competition models based on assessments of relative weed green area. An understanding of the role of various plant traits in determining early growth rate may also be useful for identifying contrasting weed strategies for establishment before canopy closure. METHODS: The response of seedling relative growth rate (RGR) to the environment was measured in outdoor sand beds in the autumn and the spring for 18 temperate annual weed species and two crops. Seedling growth was modelled using thermal time and effective day-degrees (combining the effect of temperature and radiation). The contribution of various plant traits in determining variability in RGR was investigated using regression analysis. KEY RESULTS: The effective day-degree model was more effective for describing early weed growth than thermal time. Variability in RGR measured in the autumn was largely determined by differences between the species in net assimilation rate (NAR), whereas in the spring leaf area ratio (LAR) played a larger part. There were differences between the broadleaf and grass species in the relative contribution of NAR and LAR to RGR in both seasons. RGR in the spring was negatively correlated with initial seedling size. CONCLUSIONS: The parameters derived in this study can be used to calibrate empirical models of crop yield loss based on relative weed green area to different growing seasons and assessment dates. The grass weeds, which tended to have large seeds, had a higher investment in roots in the seedling stage, potentially making them more competitive later in the season when resources become limiting.     

Spatial and temporal variation in RGR and leaf quality of a clonal riparian plant: Arundo donax

Arundo donax L. is a tall perennial reed classified as an emergent aquatic plant. In California, it has invaded riparian zones, where it acts as a transformer species. Because plant growth and leaf quality influence the effectiveness of management techniques, we sought to determine if these characters varied temporally and spatially in a northern California population of A. donax. Tissue C and N content and C:N ratio varied during the growing season. Leaf N was higher in spring and in plants that were closer to a stream. It was significantly negatively related to the clump's distance from the stream but not related to its elevation relative to the stream. Plants near the stream produced taller stems with more leaves per stem than those more distant from the stream. RGR differed across time and space. It was highest in spring prior to the appearance of flowers on a few stems that were >1 year old within the clumps. Decline in RGR as the growing season progressed coincided with the appearance of branches and flowers on stems <1 year old on a few plants within the studied population. RGR was significantly related to the N content and C:N ratio of leaves on mature stems (>1 year old). This implies that the decrease in stem growth reflected changes in nutrient availability within the entire A. donax clump and not just in the growing stems (<1 year old). These findings have implications for timing of management techniques.     

Ontogenetic patterns of leaf CO2 exchange, morphology and chemistry in Betula pendula trees

In order to explore ontogenetic variation in leaf-level physiological traits of Betula pendula trees, we measured changes in mass- (A _mass) and area-based (A _area) net photosynthesis under light-saturated conditions, mass- (RS_mass) and area-based (RS_area) leaf respiration, relative growth rate, leaf mass per area (LMA), total nonstructural carbohydrates (TNC), and macro- and micronutrient concentrations. Expanding leaves maintained high rates of A _area, but due to high growth respiration rates, net CO₂ fixation occurred only at irradiances >200 μmol photons m^–2 s^–1. We found that full structural leaf development is not a necessary prerequisite for maintaining positive CO₂ balance in young birch leaves. Maximum rates of A _area were realized in late June and early July, whereas the highest values of A _mass occurred in May and steadily declined thereafter. The maintenance respiration rate averaged ≈8 nmol CO₂ g^–1 s^–1, whereas growth respiration varied between 0 and 65 nmol CO₂ g^–1 s^–1. After reaching its lowest point in mid-June, leaf respiration increased gradually until the end of the growing season. Mass and area-based dark respiration were significantly positively correlated with LMA at stages of leaf maturity, and senescence. Concentrations of P and K decreased during leaf development and stabilized or increased during maturity, and concentrations of immobile elements such as Ca, Mn and B increased throughout the growing season. Identification of interrelations between leaf development, CO₂ exchange, TNC and leaf nutrients allowed us to define factors related to ontogenetic variation in leaf-level physiological traits and can be helpful in establishing periods appropriate for sampling birch leaves for diagnostic purposes such as assessment of plant and site productivity or effects of biotic or abiotic factors. Received: 29 December 1998 / Accepted: 26 July 1999     

Adult beetle grazing induces willow trichome defence against subsequent larval feeding

Induced increases in trichome density to grazing by herbivores have been suggested to increase plant resistance to future herbivore attack. In this paper we present results which show that plants of Salix cinerea L. respond to adult leaf beetle (Phratora vulgatissima L.) grazing by developing new leaves with an increased trichome density. The same plants are usually attacked again later in the season when the next generation of larvae hatches on the plants. The effect of the induced response was studied by comparing larval growth and feeding on newly developed leaves of plants exposed to different defoliation treatments. Larvae on plants previously exposed to adult grazing consumed less total leaf area and showed more dispersed feeding than larvae on plants protected from previous grazing. Larvae on plants exposed to mechanical defoliation responded intermediately. These results corresponded to the increased trichome density of defoliated plants. However, we found this larval response only in whole plant tests--when reared on single, excised leaves in petri dishes, larvae in all treatments behaved similarly. This discrepancy between the on-plant experiment and that in petri dishes highlights how experimental design may alter the conclusion of a study. We suggest that the induced response to adult grazing may act as a defence against subsequent larval feeding.     

川西亚高山白桦林穿透雨和茎流特征观测研究

观测了川西亚高山白桦次生林一个生长季节内的穿透雨和茎流的特征。结果表明 ,幂函数方程能较好地拟合林冠截留量、茎流量与总降雨量之间的关系 ,而线性方程能较好地拟合穿透雨量和总降雨量之间的关系 ;平均林冠截留量占总降雨量的 1 8 9% ,穿透雨量占总降雨量的 80 9% ,茎流量占总降雨量的 0 3%。