A test of the latitudinal defense hypothesis: herbivory, tannins and total phenolics in four North American tree species

It is widely believed that insect herbivory is less intense at higher latitudes, due to winter mortality which would tend to keep insect herbivores from reaching density-limitation of their populations. One prediction of this theory is that plants should tend to be better defended at lower latitudes. Here we investigated latitudinal trends in herbivory and tannins, in four species of common North American trees. Our comparisons spanned 15° of latitude in Acer rubrum, Fagus grandifolia, and Quercus alba, and 10° latitude in Liquidambar styraciflua. Sun leaves on forest edges were sampled, at phenologically equivalent times of year. Analysis revealed significant differences between populations, including those at similar latitudes, but no significant latitudinal trend in herbivory, condensed and hydrolyzable tannins, or total phenolics measured as Folin–Denis reactives in any of the four species. Our findings contradict the theory that low latitude plants are better defended, in that lower latitude populations of the four tree species showed no greater amounts of phenolics. The possible implications for community ecology are discussed. Jonathan M. Adams, Brian Rehill, and Yangjian Zhang are equal joint lead authorship on this paper.     

Phenolic-rich leaf carbon fractions differentially influence microbial respiration and plant growth

Phenolics can reduce soil nutrient availability, either indirectly by stimulating microbial nitrogen (N) immobilization or directly by enhancing physical protection within soil. Phenolic-rich plants may therefore negatively affect neighboring plant growth by restricting the N supply. We used a slow-growing, phenolic-rich alpine forb, Acomastylis rossii, to test the hypothesis that phenolic-rich carbon (C) fractions stimulate microbial population growth and reduce plant growth. We generated low-molecular-weight (LMW) fractions, tannin fractions, and total soluble C fractions from A. rossii and measured their effects on soil respiration and growth of Deschampsia caespitosa, a fast-growing, co-dominant grass. Fraction effects fell into two distinct categories: (1) fractions did not increase soil respiration and killed D. caespitosa plants, or (2) fractions stimulated soil respiration and reduced plant growth and plant N concentration while simultaneously inhibiting root growth. The LMW phenolic-rich fractions increased soil respiration and reduced plant growth more than tannins. These results suggest that phenolic compounds can inhibit root growth directly as well as indirectly affect growth by reducing pools of plant available N by stimulating soil microbes. Both mechanisms illustrate how below-ground phenolic effects may influence the growth of neighboring plants. We also examined patterns of foliar phenolic concentrations among populations of A. rossii across a natural productivity gradient (productivity was used as a proxy for competition intensity). Concentrations of some LMW phenolics increased significantly in more productive sites where A. rossii is a competitive equal with the faster growing D. caespitosa. Taken together, our results contribute important information to the growing body of evidence indicating that the quality of C moving from plants to soils can have significant effects on neighboring plant performance, potentially associated with phytoxic effects, and indirect effects on soil biogeochemistry.     

Growth,chemical responses and herbivory after simulated leaf browsing in <Emphasis Type="Italic">Combretum apiculatum</Emphasis>

Vegetative and chemical responses to simulated leaf browsing during the growth season, and their subsequent effect on herbivory, were studied on Combretum apiculatum Sonder (Combretaceae) in Botswana. Treatments (50% and 100% leaf and shoot apex removal) were performed just before the shoot growth curve levelled out, and responses recorded 3 months later, just before leaf fall. Compared to controls, defoliation treatments, removing apical dominance, reduced growth in tree height and increased shoot mortality, although the production of lateral shoots increased. At the end of the trial, there was no difference in total length of annual shoots between treatment groups. Significant refoliation occurred only after 100% defoliation. Refoliated leaves were smaller and the 100% defoliated trees had a lower final leaf biomass. Total leaf biomass production was, however, equal for all treatment groups. Refoliated leaves contained higher levels of N, lower levels of acid-detergent fibre (ADF) and total phenolics, and showed a trend towards lower levels of condensed tannins, compared to leaves on control trees. Such chemical changes may be due to either carbon stress or to younger physiological age of new leaves. In spite of the observed potential increase in food quality, we found no evidence of increased levels of insect or ungulate herbivory on refoliated leaves, which, at least for insect herbivory, may be explained by the reduction in temporal availability of leaves. We conclude that the single severe defoliation was not detrimental to C. apiculatum in the short-term, although the resource loss and induced compensatory growth may produce negative effects during subsequent growth seasons.     

Rapid leaf litter decomposition of deciduous understory shrubs and lianas mediated by mesofauna

Plant Ecology - Leaf litter decomposition rates (LDRs) of understory woody species vary substantially across species in temperate forest ecosystems. Using litter traits and LDR data for 78 shrub...     

Phylogeny and patterns of floral diversity in the genus Piper (Piperaceae)

With ～1000 species distributed pantropically, the genus Piper is one of the most diverse lineages among basal angiosperms. To rigorously address the evolution of Piper we use a phylogenetic analysis of sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA based on a worldwide sample. Sequences from a total of 51 species of Piper were aligned to yield 257 phylogenetically informative sites. A single unrooted parsimony network suggested that taxa representing major geographic areas could potentially form three monophyletic groups: Asia, the South Pacific, and the Neotropics. The position of Pothomorphe was well supported among groups of New World taxa. Simultaneous phylogenetic analysis of an expanded alignment including outgroups suggested that taxa from the South Pacific and Asia formed a monophyletic group, provisionally supporting a single origin of dioecy. Within the Neotropical sister clade, resolution was high and strong bootstrap support confirmed the monophyly of several traditionally recognized infrageneric groups (e.g., Enckea [including Arctottonia], Ottonia, Radula, Macrostachys). In contrast, some of the species representing the highly polytypic subgroup Steffensia formed a clade corresponding to the previously recognized taxon Schilleria, while others were strongly associated with several of the more specialized groups of taxa. The distribution of putatively derived inflorescence and floral character states suggested that both umbellate and solitary axillary inflorescences have multiple origins. Reduction in anther number appears to be associated with highly packaged inflorescences or with larger anther primordia per flower, trends that are consistent with the suppression of later stages of androecial development.     

成都地区10种园林灌木叶片结构与抗旱性关系研究

利用石蜡切片法和指甲油印迹法,对成都地区10种常见园林灌木叶片解剖结构的13个指标:主脉厚度、叶片厚度、上下表皮角质层厚度、上下表皮厚度、栅栏组织厚度、海绵组织厚度、下表皮气孔密度、气孔长度、气孔宽度、叶片结构紧密度、栅栏组织与海绵组织厚度比(栅海比)进行测定和分析,再用方差分析和主成分分析法选取有代表性的指标,结合隶属函数法对物种间的抗旱性进行综合评价。结果显示,13个指标的变异系数为17.40%～68.20%,在10种园林灌木间差异极显著。10种植物都属于异面叶,组织结构易于辨识(三角梅的下表皮角质层太薄,无法测出数据)。叶片结构紧密度、栅海比、气孔密度、气孔长、气孔宽、栅栏组织厚度可作为评价这10种园林灌木抗旱性的重要指标。抗旱性由强到弱依次为:红花继木、鸭脚木、红叶石楠、栀子、西洋杜鹃、海桐、山茶、雀舌黄杨、三角梅和金边大叶黄杨。其中红花继木、鸭脚木、红叶石楠和栀子属于强抗旱品种,可用于屋顶、边坡等区域种植。     

Nitrogen-addition effects on leaf traits and photosynthetic carbon gain of boreal forest understory shrubs

Boreal coniferous forests are characterized by fairly open canopies where understory vegetation is an important component of ecosystem C and N cycling. We used an ecophysiological approach to study the effects of N additions on uptake and partitioning of C and N in two dominant understory shrubs: deciduous Vaccinium myrtillus in a Picea abies stand and evergreen Vaccinium vitis-idaea in a Pinus sylvestris stand in northern Sweden. N was added to these stands for 16 and 8 years, respectively, at rates of 0, 12.5, and 50 kg N ha^?1 year^?1. N addition at the highest rate increased foliar N and chlorophyll concentrations in both understory species. Canopy cover of P. abies also increased, decreasing light availability and leaf mass per area of V. myrtillus. Among leaves of either shrub, foliar N content did not explain variation in light-saturated CO₂ exchange rates. Instead photosynthetic capacity varied with stomatal conductance possibly reflecting plant hydraulic properties and within-site variation in water availability. Moreover, likely due to increased shading under P. abies and due to water limitations in the sandy soil under P. sylvestris, individuals of the two shrubs did not increase their biomass or shift their allocation between above- and belowground parts in response to N additions. Altogether, our results indicate that the understory shrubs in these systems show little response to N additions in terms of photosynthetic physiology or growth and that changes in their performance are mostly associated with responses of the tree canopy.     

我国豆科树种结瘤调查

据国内资料报道,我国豆科树种约94属763种,已调查过结瘤状况的有含羞草亚科的123种(包括近几年来从国外引种的54种)、蝶形花亚科的211种和苏木亚科的70种,其中可以结瘤的有含羞草亚科的116种、蝶形花亚科的207种和苏木亚科的18种。含羞草亚科中的2种、蝶形花亚科中的3种均有结瘤和不结瘤的报道。     

Partitioning of 14C-labeled photosynthate to allelochemicals and primary metabolites in source and sink leaves of aspen: evidence for secondary metabolite turnover

Theories on allelochemical concentrations in plants are often based upon the relative carbon costs and benefits of multiple metabolic fractions. Tests of these theories often rely on measuring metabolite concentrations, but frequently overlook priorities in carbon partitioning. We conducted a pulse-labeling experiment to follow the partitioning of ¹⁴CO₂-labeled photosynthate into ten metabolic pools representing growth and maintenance (amino acids, organic acids, lipids plus pigments, protein, residue), defense (phenolic glycosides, methanol:water and acetone-soluble tannins/phenolics), and transport and storage (sugars and starch) in source and importing sink leaves of quaking aspen (Populus tremuloides). The peak period of ¹⁴C incorporation into sink leaves occurred at 24 h. Within 48 h of labeling, the specific radioactivity (dpm/mg dry leaf weight) of phenolic glycosides declined by over one-third in source and sink leaves. In addition, the specific radioactivity in the tannin/phenolic fraction decreased by 53% and 28% in source and sink leaves, respectively. On a percent recovery basis, sink leaves partitioned 1.7 times as much labeled photosynthate into phenolic glycosides as source leaves at peak ¹⁴C incorporation. In contrast, source leaves partitioned 1.8 times as much ¹⁴C-labeled photosynthate into tannins/phenolics as importing sink leaves. At the end of the 7-day chase period, sink leaves retained 18%, 52%, and 30% of imported ¹⁴C photosynthate, and labeled source leaves retained 15%, 66%, and 19% of in situ photosynthate in metabolic fractions representing transport and storage, growth and maintenance, and defense, respectively. Analyses of the phenolic fractions showed that total phenolics were twice as great and condensed tannins were 1.7 times greater in sink than in source leaves. The concentration of total phenolics and condensed tannins did not change in source and sink leaves during the 7-day chase period. Received: 31 July 1998 / Accepted: 8 February 1999     

我国豆科树种结瘤调查(续)

据国内资料报道,我国豆科树种约94属763种,已调查过结瘤状况的有含羞草亚科的123种(包括近几年来从国外引种的54种)、蝶形花亚科的211种和苏木亚科的70种,其中可以结瘤的有含羞草亚科的116种、蝶形花亚科的207种和苏木亚科的18种。含羞草亚科中的2种、蝶形花亚科中的3种均有结瘤和不结瘤的报道。     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Root surface/leaf area ratios in arctic dwarf shrubs

Summary Individual shoots of the shrubsLedum palustre L.,Vaccinium uliginosum L., andBetula nana L. were severed from their parent plants beneath the moss surface in an Alaskan tussock tundra. These shoots remained one year in their original position in peat moss cushions without further disturbance. After this period fine root dry weight, fine root surface area, leaf dry weight, and leaf area of these shoots were measured and compared with equivalent values from unsevered control shoots. Dry weight ratios of fine roots/leaves were similar in cut and control shoots, with the exception ofB. nana. The fine root surface/leaf area ratios showed significant differences between control and cut shoots except inL. palustre. Without tedious rootlet extractions it should be possible to estimate fine root surface area from leaf area ofL. palustre.     

Leafing patterns and leaf traits of four evergreen shrubs in the Patagonian Monte, Argentina

We assessed leafing patterns (rate, timing, and duration of leafing) and leaf traits (leaf longevity, leaf mass per area and leaf-chemistry) in four co-occurring evergreen shrubs of the genus Larrea and Chuquiraga (each having two species) in the arid Patagonian Monte of Argentina. We asked whether species with leaves well-defended against water shortage (high LMA, leaf longevity, and lignin concentration, and low N concentration) have lower leaf production, duration of the leafing period, and inter-annual variation of leafing than species with the opposite traits. We observed two distinctive leafing patterns each related to one genus. Chuquiraga species produced new leaves concentrated in a massive short leafing event (5–48 days) while new leaves of Larrea species emerged gradually (128–258 days). Observed leafing patterns were consistent with simultaneous and successive leafing types previously described for woody plants. The peak of leaf production occurred earlier in Chuquiraga species (mid September) than in Larrea species (mid October–late November). Moreover, Chuquiraga species displayed leaves with the longest leaf lifespan, while leaves of Larrea species had the lowest LMA and the highest N and soluble phenolics concentrations. We also observed that only the leaf production of Larrea species increased in humid years. We concluded that co-occurring evergreen species in the Patagonian Monte displayed different leafing patterns, which were associated with some relevant leaf traits acting as plant defenses against water stress and herbivores. Differences in leafing patterns could provide evidence of ecological differentiation among coexisting species of the same life form.     

Adventitious shoot regeneration from root, internode, petiole and leaf explants of Piper colubrinum Link.

A single-step, high-frequency regeneration pro-tocol has been standardised for Phytophthora-resistant wild pepper, Piper colubrinum (Link) using root, internode, leaf and petiole explants derived from in vitro plantlets. The effect of BA on shoot-bud induction and elongation was assessed by supplementing half-strength MS medium (macronutrients at half the concentration) with different concentrations of BA, i.e. 0.2–10 mg l^–1 in induction media and 0, 0.2 and 0.5 mg l^–1 in subculture media. The interaction between culture period and BA concentration was studied by culturing the explants for 8, 15 and 30 days before the first subculture. The elongated shoots were rooted directly in soil and hardened in the greenhouse. The developed protocol would be useful in marker-assisted asymmetric hybridisation programmes involving wild-type Piper colubrinum and the cultivated species P. nigrum. Received: 4 August 1997 / Revision received: 30 January 1998 / Accepted: 12 February 1998     

Drought constraints on leaf gas exchange by Miconia ciliata (Melastomataceae) in the understory of an eastern Amazonian regrowth forest stand

Analyses of the effects of drought stress on Amazonian regrowth stands are lacking. We measured leaf gas exchange and leaf water potential of Miconia ciliata (Melastomataceae) in a dry-season irrigation experiment in 14-yr-old regrowth. In the dry season, irrigated plants maintained significantly higher leaf water potentials, photosynthetic capacity at light saturation (A(max)), stomatal conductance (g(s)), internal CO(2) concentration (C(i)), and lower A(max)/g(s) than control plants. The degree of dry-season down-regulation of control plant A(max), along with its fast recovery following rain, reveals the importance of occasional dry-season rains to the carbon budget of M. ciliata. During the wet season, we observed higher A(max) for control plants than for plants that had been irrigated during the dry season. We hypothesize that reduced drought constraints on photosynthesis of irrigated plants advanced the flowering and fruiting phenology of irrigated plants into the dry season. Flowers and fruits of control plants developed later, during the wet season, potentially stimulating a compensatory reproductive photosynthesis response in nearby leaves. The relative drought intolerance of M. ciliata may be a deciding factor in its ability to survive through the dynamic successional development of the regrowth stand studied.     

Ecophysiology of exotic and native shrubs in Southern Wisconsin

Summary We compared seasonal trends in photosynthesis of two naturalized exotic shrubs (Rhamnus cathartica and Lonicera X bella) and two native shrubs (Cornus racemosa and Prunus serotina) in open and understory habitats in southern Wisconsin. We examined the relationships between resource availability and leaf photosynthetic performance in these four species. All four species had similar relationships between leaf nitrogen (N) content and photosynthetic rate, but the species differed in absolute leaf N content and therefore in photosynthetic rates. Maximum daily photosynthetic rates of all species were significantly correlated with leaf N content in the open habitat, but not in the understory, where low light availability was the major limitation to photosynthesis. Extended leaf longevity was important in the forest understory because it allowed shrubs to take advantage of high light availability at times when the overstory canopy was leafless. Early leaf emergence was more important than late senescence: from 27% to 35% of the annual carbon gain of P. serotina, R. cathartica, and L. X bella occurred prior to leaf emergence of C. racemosa, the species with the shortest leaf life span. Extended leaf longevity of exotic shrubs may help explain their persistence in the understory habitat, but it contributed relatively less to their annual carbon gain in the open habitat.     

祁连山典型灌丛降雨截留特征

基于2010年6月至10月的野外试验数据,研究了祁连山金露梅、高山柳、沙棘和鬼箭锦鸡儿灌丛降雨截留特征,分析了降雨量和雨强对灌丛降雨截留过程的影响。结果表明:试验期间共降雨298.6 mm,在降雨量<2.1 mm时,降雨被全部截留,实际发生穿透和茎流的降雨为283.1 mm;金露梅灌丛穿透雨量、茎流量和截留量分别为175.8 mm(62.0%)、9.5 mm(3.4%)和62.0 mm(34.6%),高山柳为179.8 mm(63.5%)、9.1 mm(3.2%)和63.5 mm(33.3%),沙棘分别为148.1mm(52.3%)、22.5mm(8.0%)和52.3 mm(39.7%),鬼箭锦鸡儿分别为170.4 mm(60.2%)、11.8 mm(4.2%)和60.2 mm(35.6%);灌丛穿透雨量、茎流量和截留量均与降雨量呈显著线性正相关(P<0.001);穿透率、茎流率和截留率与降雨量呈指数函数关系(P<0.05);平均雨强与截留率关系以指数函数拟合最好(P<0.05)。在降雨性质相同的情况下,植被形态特征是影响灌丛降雨截留的重要因素。     

Role of defense enzymes and phenolics in resistance of wheat crop (Triticum aestivum L.) towards aphid complex

Herbivory damage leads to induction of rapid signals and responses in plants such as oxidative burst, accumulation of secondary metabolites and defensive proteins. Response of various defensive enzymes and secondary metabolites in flag leaf samples of six bread wheat varieties against aphid feeding was investigated. Six bread wheat varieties, namely PBW 621 and HD 2967 (timely sown irrigated), PBW 590 and PBW 658 (late sown irrigated), and PBW 644 and PBW 660 (timely sown rainfed) were grown under the aphid infested and uninfested conditions and were sampled at a regular interval to analyze the biochemical changes caused by aphid feeding. A tremendous increase in the overall activity of various enzymes namely superoxide dismutase, glutathione reductase, phenylalanine ammonia lyase and polyphenol oxidase was observed, all of which play an important role in plants defense towards aphid feeding. Each wheat genotype showed an overall difference in their defensive activity towards aphid feeding. However, certain genotypes under different conditions showed significantly less susceptibility towards aphid damage.

Abbreviations: GR: glutathione reductase; HPR: host plant resistance; PAL: phenylalanine ammonia lyase; PPO: poly phenol oxidase; POD: peroxidase; SOD: superoxide dismutase