Consequences of herbivory in the mountain birch (Betula pubescens ssp tortuosa): importance of the functional organization of the tree

Summary Three types of experiments indicate that the functional organization of the mountain birch may influence the ways in which the tree responds to simulated or natural herbivory. The first experiment showed that herbivory to both short and long shoot leaves affects plant development but, because growth largely proceeds by resources of the previous year, is manifested only in the year following the damage. The second experiment showed that even partial damage to a single long shoot leaf caused the axillary bud of that leaf to produce a shorter shoot the next year. Therefore, the value of a leaf depends also on the organ which it is subtending. In the third experiment we manipulated the apical dominance of shoots in ramets and caused improvement to leaf quality in extant shoots. Ramets within a tree responded individually, probably mediated by disturbance of the hormonal control because removal of apical buds elicited the response although removal of the same number of basal buds did not. Induced amelioration is a different response to induced resistance. The two responses are triggered by different cues and may occur in the same plant. By altering hormonal balance of shoots it is potentially possible for herbivores to induce amelioration of food quality. The ways in which herbivory is simulated may explain variability of results obtained when herbivory-induced responses in plants have been studied.     

Natural hybridisation in birch: triploid hybrids between Betula nana and B. pubescens

Hybridisation between diploid (2n=28) dwarf birch Betula nana L. and tetraploid (2n=56) downy birch B. pubescens Ehrh. has occurred in natural populations in Iceland. About 10% of birch plants randomly collected are triploid (2n=42) hybrids. Ribosomal gene mapping on chromosomes and genomic in situ hybridisation confirms the hybridity. However, the triploid hybrids are not morphologically distinct, i.e. they are not different from diploid and tetraploid birch plants that have intermediate morphology. The triploid hybrids have evidently played an important role in driving bi-directional gene flow between these two species. This paper reviews the extent of interspecific hybridisation in selected birch woodland populations and discusses the significance of natural hybridisation and introgression in birch.     

Response of leaf surfaces and gas exchange to wind stress and acid mist in birch (Betula pubescens)

 This research demonstrates that a leaf’s response to acid mist is dependent on the integrity of the leaf cuticle and that significant differences in the structural and physiological disturbances in leaves can be attributable to different types of wind action. Betula pubescens Ehrh. plants were located at adjacent, but contrasting, sites to create different wind treatments: (i) direct wind action, (ii) indirect wind action and (iii) shelter from wind action (control). In combination with the wind treatments, acidic (pHs 5 and 3) or neutral (pH 7) mists were applied weekly. Wind action significantly increased visible leaf injury, microscopic cuticular lesions and cuticular conductance (g _c ), but reduced photosynthetic rate (P _N ) and stomatal conductance (g _s ) compared to shelter. Wind action combined with acid mist was more injurious than wind action alone, but leaves sheltered from wind action were highly resistant to the damaging effects of acid mist. Direct wind action combined with pH 3 mist resulted in the highest values of g _c and the greatest number of cuticular lesions. By contrast, indirect wind action combined with pH 3 mist induced most visible injury, but relatively low values of g _c and few microscopic cuticular lesions. Acid mist reduced P _N only when leaves had been damaged by wind action. Higher values of g _c were associated both with increases in the area of visible leaf injury and with the number of cuticular lesions. Compensatory increase in P _N of healthy tissue was evident in leaves exposed to combinations of wind action and acid mist. Received: 10 November 1997 / Accepted: 6 March 1998     

Recent invasion of the mountain birch Betula pubescens ssp. tortuosa above the treeline due to climate change: genetic and ecological study in northern Sweden

Mountain birch, Betula pubescens ssp. tortuosa, forms the treeline in northern Sweden. A recent shift in the range of the species associated with an elevation of the treeline is commonly attributed to climate warming. Using microsatellite markers, we explored the genetic structure of populations along an altitudinal gradient close to the treeline. Low genetic differentiation was found between populations, whereas high genetic diversity was maintained within populations. High level of gene flow compensated for possible losses of genetic diversity at higher elevations and dissipated the founding effect of newly established populations above the treeline. Spatial autocorrelation analysis showed low spatial genetic structure within populations because of extensive gene flow. At the treeline, significant genetic structure within the juvenile age class at small distances did not persist in the adult age class, indicating recent expansion of young recruits due to the warming of the climate. Finally, seedling performance above the treeline was positively correlated with parameters related to temperature. These data confirm the high migration potential of the species in response to fluctuating environmental conditions and indicate that it is now invading higher altitudes due to the recent warming of the climate.     

Sexual and asexual reproduction in geographically separated populations of the fissiparous asteroid Coscinasterias calamaria (Gray)

Sexual reproduction and asexual reproduction by fission were studied in four populations of Coscinasterias calamaria (Gray), two in Otago Harbour in the South Island of New Zealand and two in the North Island near Auckland. The annual reproductive cycle in both islands of New Zealand is clearly defined with a spawning season between November and January. In both sites the pyloric caeca index was approximately inverse to the gonad index cycle as found in other forcipulate asteroids. There are substantial differences in the sex ratios of mature starfish at each site studied, with 1:1 ratios in two populations, one population heavily biased towards females and the fourth consisting almost entirely of males. Morphometric variation in arm number due to splitting was studied and the frequency of splitting varied considerably between the four populations. Generally speaking sublittoral C. calamaria divide less frequently than intertidal starfish and populations in which food is less abundant or of poorer quality are more fissiparous and put less energy into sexual reproduction, than populations with plentiful readily available food in the form of mussels. The significance of the different reproductive patterns in C. calamaria is discussed.     

浙江丽水中国石龙子的食性、两性异形和雌性繁殖

丽水分布的中国石龙子（Ｅｕｍｅｃｅｓ ｃｈｉｎｅｎｓｉｓ）摄入的食物均为无脊椎动物,分别属于环节、软体和节肢动物,涉及３０余科,成体和幼体的食物生态位宽度分别为７．２６和６．６９,成体的幼体的食物生态们重叠度为０．５９。性成熟雄性个体大于雌体。成雄和幼体的头长和头宽随体长ＳＶＬ的增长速率大于成雌,成雄头长随ＳＶＬ的增长速度显著大于幼体,成雌和幼体的头长随ＳＶＬ的增长速率无显著差异。成雄头部大于成雌     

Sexual reproduction in Daphnia pulex (Crustacea: Cladocera): observations on male mating behaviour and avoidance of inbreeding

1.  Mating behaviour in Daphnia appears to rely on random contact between males and sexual females rather than diffusible pheromones. Males may be able to discriminate sexually receptive females from females in other developmental stages and increase their mating efficiency. Males may also use chemical signals to avoid mating with females from the same clone and avoid the severe inbreeding depression that has been documented for intraclonal mating. The present study used experiments to test for the avoidance of intraclonal mating and assess male mating efficiency in D. pulex .
2.  Three clones were examined for the avoidance of intraclonal mating by providing males with an opportunity to mate with females of the same or two different clones. The proportion of intraclonal matings did not differ from the proportion of interclonal matings, suggesting that D. pulex males do not use kin discrimination to avoid mating with females from the same clone.
3.  The proportion of mated females decreased with increasing numbers and density of sexual females when exposed to a single male. This observation suggests that a male spends more time pursuing and copulating with sexually receptive females than non-receptive females and there is insufficient time to mate with all sexual females. The decrease in proportion of females mated could also be the result of sperm depletion in the male. Sperm depletion is unlikely to occur in nature because sexually receptive females are much rarer than in the experimental conditions.     

Dependence of amino acid composition upon nitrogen availability in birch (Betula pendula)

Small birch plants ( Betula pendula Roth .) were grown at different rates of exponentially increasing nitrogen supply. This resulted in plants with different relative growth rates and different internal nitrogen concentrations. Within a nitrogen treatment, both of these variables remained constant with time.
Free amino acids were measured in leaves and roots of the seedlings at two different harvests. At greater nitrogen supply, higher concentrations of total amino acid nitrogen were found in roots and leaves. The ratio of amino acid nitrogen to total nitrogen was low albeit greater at higher nitrogen supply. Higher concentrations of amino acid nitrogen were mainly due to high concentrations of citrulline, glutamine, γ-aminobuitric acid and arginine.
Greater leaf concentrations of amino acid nitrogen at higher nitrogen supply may be related lo increased concentrations in the xylem sap and/or may be indicative of small excesses of nitrogen with respect to current nitrogen usage in protein synthesis.     

Ecological role of reindeer summer browsing in the mountain birch (<Emphasis Type="Italic">Betula pubescens</Emphasis> ssp. <Emphasis Type="Italic">czerepanovii</Emphasis>) forests: effects on plant defense,litter decomposition,and soil nutrient cycling

Mammalian herbivores commonly alter the concentrations of secondary compounds in plants and, by this mechanism, have indirect effects on litter decomposition and soil carbon and nutrient cycling. In northernmost Fennoscandia, the subarctic mountain birch (Betula pubescens ssp. czerepanovii) forests are important pasture for the semidomestic reindeer (Rangifer tarandus). In the summer ranges, mountain birches are intensively browsed, whereas in the winter ranges, reindeer feed on ground lichens, and the mountain birches remain intact. We analyzed the effect of summer browsing on the concentrations of secondary substances, litter decomposition, and soil nutrient pools in areas that had been separated as summer or winter ranges for at least 20 years, and we predicted that summer browsing may reduce levels of secondary compounds in the mountain birch and, by this mechanism, have an indirect effect on the decomposition of mountain birch leaf litter and soil nutrient cycling. The effect of browsing on the concentration of secondary substances in the mountain birch leaves varied between different years and management districts, but in some cases, the concentration of condensed tannins was lower in the summer than in the winter ranges. In a reciprocal litter decomposition trial, both litter origin and emplacement significantly affected the litter decomposition rate. Decomposition rates were faster for the litter originating from and placed into the summer range. Soil inorganic nitrogen (N) concentrations were higher in the summer than in the winter ranges, which indicates that reindeer summer browsing may enhance the soil nutrient cycling. There was a tight inverse relationship between soil N and foliar tannin concentrations in the winter range but not in the summer range. This suggests that in these strongly nutrient-limited ecosystems, soil N availability regulates the patterns of resource allocation to condensed tannins in the absence but not in the presence of browsing.     

Reproduction of an early-flowering Mediterranean mountain narrow endemic (Armeria caespitosa) in a contracting mountain island

Reproduction at population lower edges is important for plant species persistence, especially in populations on contracting high-mountain islands. In this context, the ability of plants to reproduce in different microhabitats seems to be important to guarantee seed production in stressful environments, such as Mediterranean high mountains. We hypothesised that the warmer and drier conditions at the lower edge would be deleterious for the reproduction of Armeria caespitosa, an early-flowering plant. In addition, reproductive plasticity along this mountain gradient may also be microhabitat-dependent. We studied factors affecting the reproductive success of A. caespitosa , an endemic of the Spanish Sistema Central. We considered a complex set of predictors, including phenology, plant size and environmental factors at different scales using generalised estimating equations and generalised linear models. Microhabitat, together with the position in the altitudinal gradient and inter-annual variability affected the reproduction of A. caespitosa . In addition, individuals with longer flowering periods (duration of flowering) had significantly lower fruit set and a higher number of unviable seeds; delayed flowering peaks favoured the production of both viable and unviable fruits. Microhabitat variability over an altitudinal range is relevant for the reproduction of A. caespitosa, and is more important at the lower edge of the altitudinal range, where the species faces the most adverse conditions. In addition, the ability to reproduce in different microhabitats might increase the chances of the species to cope with environmental uncertainties under on-going climate warming. Finally, reproduction of this early-flowering plant is constrained by summer drought, which might shape its reproductive phenology.     

Effects of ultraviolet-B radiation on growth, mycorrhizas and mineral nutrition of silver birch (Betula pendula Roth) seedlings grown in low-nutrient conditions

The effects of increased ultraviolet‐B (UV‐B) radiation on the growth, mycorrhizas and mineral nutrition of silver birch (Betula pendula Roth) seedlings were studied in greenhouse conditions. Seedlings—planted in a birch‐forest top soil and sand substrate—were grown without additional nutrient supply. Ultraviolet treatment started immediately after the seedlings emerged and the daily integrated biologically effective UV‐B irradiance on the UV‐B‐treated plants was equivalent to a 25% depletion of stratospheric ozone under clear sky conditions. Visible symptoms of UV‐B damage or nutrient deficiency were not observed throughout the experiment. Seedling height and dry weight (DW) (measured after 58 days and 76 days of treatment) were not affected by increased UV‐B. However, a significant shift in DW allocation toward roots resulted in a lower shoot/root ratio and leaf area ratio in UV‐B‐treated plants compared to control plants. At the first harvest (after 58 days of treatment), the percentage of various mycorrhizal morphotypes and the number of short roots per unit of root length or weight were not affected by increased UV‐B despite significantly increased DW allocation toward roots. Subtle reduction in the allocation of nitrogen (N) to leaves and increased allocation of phosphorus (P) to roots may suggest cumulative effects that could affect the plant performance over the long‐term.     

Estimation of canopy density and irradiance in 20- to 40-year-old birch stands (Betula pubescens Ehrh. andBetula pendula Roth)

Three different experiments in birch (Betula pubescens Ehrh. andBetula pendula Roth) stands were conducted during summer 1991: an evaluation of the accuracy of the vertical tube method measuring crown free projection (CFP) in four birch stands; a study of relative irradiance (RI; photosynthetically active radiation, PAR), in birch stands differing in stem density and basal area; and an estimation of the CFP in different types of birch stands and the correlation between RI and CFP. The results indicate that the number of observation points well distributed over an area of 0.1 ha can vary between 50 and 200 to achieve sufficient accuracy. There were no significant differences in the CFP estimates made on the same plot by different observers including an untrained observer. CFP was related to basal area, stem number and diameter sum in a non-linear regression, CFP = A^*e^-BX. The correlations between RI and basal area, density and diameter sum, respectively, were 0.54, 0.57 and 0.67 for PAR. The coefficients of determination for RI (PAR) and CFP were high (0.86).     

Sexual generation and overwintering of the peach leaf curling aphid Brachycaudus helichrysi (Kalt.) in Himachal Pradesh, India

Autumn migrants preferred to settle on the dorsal leaf surface of peach with gynoparae being more numerous than alate males. These return migrants on peach were similar to those described by others except for a comparatively smaller processus terminalis. Among the oviparae with metatibial pseudosensoria. most (62.5%) had six antennal subdivisions; 25% possessed five antennal subdivisions and half of these were parasitised. Oviparae were bigger than those previously described on plum/damson but their processus terminalis was comparatively shorter. Ultimate instar nymphs were without pseudosensoria and had 5–seg-mented antennae, the VI segment being cleaved from the meriston during the final moult. Alate males mated with the oviparae, mating duration being 3 to 6 min. Oviparae contained, at the most, two well developed eggs at one time, and normally deposited eggs singly during November-December. Most eggs hatched within 20–28 days but those laid in December during colder weather took up to 50 days to hatch. By midJanuary most eggs had hatched. Fundatrix developmental rate was dependent on sap-flow. Overwintering occurred as the fundatrix.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

Sexual swelling in mandrills (Mandrillus sphinx): a test of the reliable indicator hypothesis

Various hypotheses have been proposed to account for the functionof sexual swellings in female primates, but few empirical dataexist to test predictions arising from these hypotheses. Controversyhas recently arisen over a field study that appeared to supportthe predictions of the reliable indicator hypothesis. This hypothesisproposes that females compete for males or matings, that differencesin swelling size between females reliably advertise female quality,and that males use swelling characteristics to differentiallyallocate mating effort to females with certain swelling characteristics,hence to females of higher quality. To provide an independenttest of this hypothesis, we collected data concerning the sizeand coloration of 40 sexual swellings for 29 semi-free-rangingfemale mandrills, varying in age and parity, along with dataconcerning the behavior of males toward the females, and comparedthese with the long-term reproductive history of the females.We examined the following predictions: (1) swelling characteristicsare consistent across subsequent cycles for individual females,(2) swelling characteristics indicate aspects of female reproductivequality, and (3) males prefer to mate with females that showparticular swelling characteristics. Our results support prediction1; we found little change in swelling characteristics acrossswellings for individual females. However, we found no significantrelationships between female reproductive history and swellingcharacteristics and, thus, no support for prediction 2. Finally,we found only limited support for prediction 3; females withlarger (wider) sexual swellings were more likely to have a spermplug when maximally swollen. However, male mate-guarding wasnot significantly related to female swelling characteristics.Furthermore, in situations in which more than one female wasmaximally swollen, the alpha male (who has "free" choice) didnot show the most interest in the female with the largest swelling.We conclude that the reliable indicator hypothesis does notexplain variation in sexual swellings in female mandrills.     

Transpiration rates of the understory annual Impatiens capensis (Balsaminaceae) in response to the autumnal changes in canopy leaf area

Leaf transpiration rates of Impatiens capensis were measured beneath a broadleaved deciduous forest canopy over successive growing seasons using a steady-state porometer. The transpiration measurements, which continued into early autumn, provided a framework for assessing whether I. capensis exhibits stomatal opening in response to the autumnal increase in available direct-beam radiation reaching the forest floor. The deciduous canopy LAI (leaf area index) decreased from a growing season maximum of 3.94 m² m⁻², while the understory I. capensis population located along a stream channel maintained LAI values ranging from 0.58 to 1.05 m² m⁻² late into the growing season. Late morning and early afternoon leaf transpiration rates during the months of June and July averaged about 8 μg cm⁻² s⁻¹, with a mean stomatal conductance of 0.5 cm s⁻¹. In August, leaf transpiration averaged almost 12 μg cm⁻² s⁻¹, with stomatal conductance exceeding 1.5 cm s⁻¹. However, beginning in early to mid-September, before canopy leaf-fall, the persistent green leaves of I. capensis exhibited a sharp decline in transpiration, possibly a result of decreasing vapor pressure deficits or non-lethal physiological damage induced by cold stress. This physiological decline offsets any advantage that could have been gained by the increased exposure to direct-beam radiation after canopy leaf-fall in mid-October. Although green leaf area and seed-bearing capsules may persist until the first frost in October or early November, there is no evidence of stomatal opening suggestive of carbon assimilation for enhanced seed development during this early autumn period. We conclude that the persistent green leaf area of I. capensis fails to exploit the increase in available direct-beam radiation in the final stage of its life cycle.