The effect of forest management operations on population performance of Vaccinium myrtillus on a landscape-scale

We have studied the variation in average performance of Vaccinium myrtillus L. among local populations within an intensively managed boreal forest landscape in southeast Norway. Our aim was to identify whether forest disturbance history was an important factor controlling the variation in average plant performance. We used three measures of forest maturity (maturity class, tree biomass and number of trees) as variables reflecting disturbance history. These variables were used in our analysis together with measured environmental conditions (soil moisture, pH, nitrogen, and elevation) a priori known to affect plant performance. To quantify plant performance, we used multiple measures of the local V. myrtillus populations, including percentage cover, average ramet age, height, diameter, biomass, growth, and sexual reproduction (number of fruits and number of mature seeds per fruit). Most of the explained variation in our performance variables could be related to our environmental variables, but disturbance history was also an important determinant of the performance of V. myrtillus. All performance variables were higher in younger, less mature stands, except ramet age, which was highest in mature stands. The increase in performance after clear cutting indicates that this period is of high importance for the population dynamics and persistence of V. myrtillus in intensively managed boreal forest landscapes.     

Pollen limitation in three sympatric species ofVaccinium (Ericaceae) in the Upper Ardennes, Belgium

The reproductive success of three co-flowering species ofVaccinium (V. myrtillus, V. vitis-idaea andV. uliginosum) was studied in one heathland of the Upper Ardennes, Belgium, during three years (1988–1990). The purpose was to examine whether pollen limitation, flower position and flowering phenology may influence patterns of fruit set in these three sympatric species. I quantified fruit and seed set following supplementary hand-pollinations and compared this to natural fruit set. On the same plants, I also quantified fruit and seed set in relation to the spatial position of the flowers on the ramet and their temporal sequence of blooming. Hand-pollination had no significant effect on fruit set inV. vitis-idaea andV. uliginosum, but significantly increased seed number per fruit in 1989. InV. myrtillus both fruit and seed set were increased by supplementary pollination, but significantly in only one year. Analyses of position effects revealed that the fruits in the lower positions in the ramet did not mature preferentially and did not contain more seeds inV. uliginosum and inV. vitis-idaea. Flowering phenology also had no significant effect for these species. InV. myrtillus hand-pollinated flowers showed a seasonal decline in seed number, and control (naturally pollinated) flowers showed a seasonal increase in fruit set. Fruit and seed set appear to be pollen-limited rather than resource-limited inV. uliginosum. In the two other species, unfavorable weather (frosts) could be a more important cause of low fruit and seed set.     

Rates of cross-pollination inLycopersicon pimpinellifolium: Impact of genetic variation in floral characters

Cross-pollination was tested between different genotypes ofLycopersicon pimpinellifolium in an experimental plot situated in the southerly range of the species. Rates of cross-pollination were found to be highly correlated with flower size and were related to a lesser extent with degree of stigma exsertion. Comparisons of fertility suggest that exsertion of 1.5 mm or more from the anther tube orifice interferes with automatic self-pollination, thereby increasing rates of cross-pollination. A single species ofExomalopsis (Anthophoridae), observed to visit all tested genotypes ofL. pimpinellifolium, was responsible for most of the pollination. Other species of solitary bees and other bee families participated in cross-pollination.     

Reproductive biology of twoPrunus species (Rosaceae) in the Northwest Iberian Peninsula

We studied the reproductive biology ofPrunus spinosa andPrunus mahaleb (Prunoideae, Rosaceae) in the northwest Iberian Peninsula. The two species flowered at the same time (peaking on March 9 and 11, respectively in 1990) but differ significantly in their fruit maturation times. Nectar volume peaked in the early morning in both species, and was ten times greater inP. spinosa than inP. mahaleb. Neither species shows apomixis, nor does fruit-set occur if pollinators are excluded. In both species self-pollination resulted in fewer fruits than open pollination. The principal pollinators belong to theApidae family (79% and 63% of visits toP. spinosa andP. mahaleb, respectively). Results are compared with those for other rosaceous plants with fleshy fruits.     

Self pollination and resource availability affect ovule abortion inCassia fasciculata (Caesalpiniaceae)

We examined the effects of pollen source and resource availability on ovule abortion in the annual legumeCassia fasciculata. Pollen source was controlled by hand-pollinating flowers with cross- or self-pollen. Resource availability to developing fruits was controlled by adjusting fruit loads (heavy versus light) on each plant and exposing plants to different photoperiod cycles (16 vs 12 h of light; short days favor fruit growth at the expense of vegetative growth). In mature fruits the proportion of ovules expanding (showing some development over virgin ovules) ranged from 89–95% and did not increase with resource availability, suggesting that unexpanded ovules were either unfertilized or obligately aborted shortly after fertilization. The proportion of expanded ovules maturing in mature fruits was near 97% for both self- and crosspollinations in the treatment with highest resource availability (light load, short days) and lower in the remaining treatments, where self-pollination resulted in up to 9% lower seed maturation than cross-pollination. In the latter three treatments a pollen source effect was dependent upon the maternal plant; in some plants selfing increased abortion and in others it did not. Collectively, the results suggest that (1) both pollen source and resource availability affect ovule abortion, (2) at least some abortion is facultative, and (3) when resources are limited, self-pollination increases abortion in some but not all maternal plants.     

Effect of N6-isopentenyladenine concentration on growth initiation in vitro and rooting of bilberry and lingonberry microshoots

Buds and shoot tips of wild bilberry (Vaccinium myrtillus L.) and lingonberry (V. vitis-idaea L.) plants were cultured on a modified MS medium containing N⁶-isopentenyladenine (2iP), 9.8–78.4 μM, in order to study the effect of the 2iP-concentration on the initiation of growth. The experiment was first performed in the autumn and repeated in the spring to determine the influence of season on growth initiation. To optimise rooting, three different rooting treatments were tested for the bilberry and lingonberry microshoots. Shoots were rooted either in vitro with 0.49 μM IBA (indole-3-butyric acid) or ex vitro, incubating microshoots in 2.07 mM KIBA-solution (potassium salt of IBA) before planting, or microshoots were planted directly on peat without exogenous auxin. The best 2iP concentration for the initiation of the growth for bilberry was 49.2 μM and for lingonberry 24.6 μM. It was observed that increasing the 2iP concentration at the growth initiation stage increased the number of brownish explants both in bilberry and in lingonberry microcultures. Spring was a considerably better time than autumn for the initiation of new growth, for both species. The results of the rooting test showed that the KIBA-treatment before planting on peat increases rooting efficiency in both bilberry and lingonberry. This revised version was published online in June 2006 with corrections to the Cover Date.     

Quantitatively determined self-incompatibility

Summary It has been claimed that Borage (Borago officInalis L.) has a multifactorial self-incompatibility system. Such systems may have a high level of ineffective pollination, and we show that this is the case in borage. The ranking of seed set from highest to lowest is as follows: bee-pollination; natural pollination in the absence of bees; artificial cross-pollination between unrelated plants; artificial cross-pollination between related plants; artificial self-pollination. In diallel crosses, significant parental effects were detected but no consistent patterns of seed set, which suggest a simple self-incompatibility system, were detected. The level of outcrossing with natural pollination was very variable but greater than 50%. Thus, there appears to be no straightforward self-incompatibility system in borage.     

Pollination and self-interference in Nothofagus

Interference between male and female functions within a monoecious plant may hinder crossing and decrease seed set. We assessed the probability of self-pollination and the effect of self-pollination on cross-pollination for two self-incompatible species: Nothofagus obliqua and N. nervosa. The probability of self-pollination was studied by tracking the phenologies of staminate and pistillate flowers, including an analysis of stigmatic receptivity. Pure and mixed pollinations were performed in order to evaluate the effect of self-pollination upon cross-pollination. Phenological observations suggest that self-pollination is highly likely in both species. Compared to pure cross-pollination, the application of self-pollination prior to cross-pollination resulted in lower numbers of germinated pollen grains for both species, and also in a lower production of viable seeds in N. nervosa. The low proportion of viable seeds often observed in natural populations of N. obliqua and N. nervosa may be related to self-pollination.     

Biomass distribution of two subalpine dwarf‐shrubs in relation to soil moisture and nutrient content

Question: Do soil water content and/or soil nitrogen (N) content and/or soil phosphorus (P) content affect the biomass of Vaccinium myrtillus and V. vitis‐idaea in a sub‐alpine heath? Location: Dolomites, northern Italy, 1800 m a.s.l. Methods: We determined above‐ground and below‐ground biomass of the shrubs at three sites, each on a different substrate type. At each site, we determined soil N‐ and P‐contents. We also determined leaf water potential (Psi;₁), N‐ and P‐concentrations in plant tissues and litter, as well as δ¹³C and δ¹⁵N in mature leaves. Results: V. myrtillus biomass was highest at the silicate site, V. vitis‐idaea biomass was highest at the carbonate site. Both shrubs had low biomass at the peat site, possibly due to a toxic effect of waterlogging in wet soils. For both species, pre‐dawn Psi;₁ indicated optimal hydration and midday Psi;₁ did not show any sign of water stress. Water use efficiency (WUE) did not differ among sites for any species. Whole‐plant nutrient concentrations showed that, with increasing biomass, N was diluted in V. myrtillus tissues while P was diluted in V. vitis‐idaea tissues. Foliar N‐concentration was higher overall for V. myrtillus. Foliar P‐concentration in V. myrtillus peaked at the silicate site. Foliar N : P ratios suggested that V. myrtillus was primarily P‐limited and V. vitis‐idaea primarily N‐limited. Conclusions: Water content affected the distribution of the two shrubs in a similar way, higher P‐availability in the soil enhanced V. myrtillus rather than V. vitis‐idaea.     

Seasonal changes in plasma membrane H<Superscript>+</Superscript>-ATPase activity of <Emphasis Type="Italic">Vaccinium myrtillus</Emphasis> (L.) and <Emphasis Type="Italic">Pinus sylvestris</Emphasis> (L.)

Seasonal changes in vanadate sensitive plasma membrane H⁺-ATPase activity of bilberry (Vaccinium myrtillus L.) and Scots pine (Pinus sylvestris L.) were studied in a period from February to August in northern Finland. The plasma membrane isolation was performed by sucrose gradient centrifugation, and the H⁺-ATPase activity was assayed by spectrophotometrical determination of released inorganic phosphate. The studied species showed seasonal changes from high winter to low spring activity, indicating probable physiological changes between hardened and dehardened tissue. ATPase activity of bilberry peaked up at the beginning of the growth period, obviously due to active phloem loading of photosynthates.     

Growth and reproduction of the alpine grasshopper <Emphasis Type="Italic">Miramella alpina</Emphasis> feeding on CO<Subscript>2</Subscript>-enriched dwarf shrubs at treeline

The consequences for plant-insect interactions of atmospheric changes in alpine ecosystems are not well understood. Here, we tested the effects of elevated CO₂ on leaf quality in two dwarf shrub species (Vaccinium myrtillus and V. uliginosum) and the response of the alpine grasshopper (Miramella alpina) feeding on these plants in a field experiment at the alpine treeline (2,180 m a.s.l.) in Davos, Switzerland. Relative growth rates (RGR) of M. alpina nymphs were lower when they were feeding on V. myrtillus compared to V. uliginosum, and were affected by elevated CO₂ depending on plant species and nymph developmental stage. Changes in RGR correlated with CO₂-induced changes in leaf water, nitrogen, and starch concentrations. Elevated CO₂ resulted in reduced female adult weight irrespective of plant species, and prolonged development time on V. uliginosum only, but there were no significant differences in nymphal mortality. Newly molted adults of M. alpina produced lighter eggs and less secretion (serving as egg protection) under elevated CO₂. When grasshoppers had a choice among four different plant species grown either under ambient or elevated CO₂, V. myrtillus and V. uliginosum consumption increased under elevated CO₂ in females while it decreased in males compared to ambient CO₂-grown leaves. Our findings suggest that rising atmospheric CO₂ distinctly affects leaf chemistry in two important dwarf shrub species at the alpine treeline, leading to changes in feeding behavior, growth, and reproduction of the most important insect herbivore in this system. Changes in plant-grasshopper interactions might have significant long-term impacts on herbivore pressure, community dynamics and ecosystem stability in the alpine treeline ecotone.     

Reproductive biology ofAsphodelus aestivus (Asphodelaceae)

InAsphodelus aestivus the large number of showy flowers opened per inflorescence, high nectar and pollen production, and absence of automatic self-pollination, so that insect visitation is necessary for pollination, indicate that cross-fertilization is favoured. LargeHymenoptera are the main pollinators. Hand self-pollinations result in some degree of fruit-set, but cross-pollination also yield low fruit- and seed-set. The perennial habit and polyploid status are in agreement with the breeding system of this species, with sexual reproduction being rather inefficient and active vegetative propagation.     

紫花含笑传粉生物学初探

紫花含笑(Michelia crassipes)花色艳丽、花香浓郁,为解释其在自然条件下结实率低的原因和判断最佳杂交时期,该研究从传粉生物学入手,通过观察紫花含笑开花动态,运用异交指数(OCI)、花粉胚珠比(P/O)、花粉和柱头活性、人工授粉试验等方法对其繁育系统进行了检测,并记录访花昆虫的种类和行为。结果表明:(1)紫花含笑花期在4—5月,持续50 d左右,单花花期4~5 d。(2)雌雄同花,雌蕊先成熟,雄蕊空间位置高于雌蕊。(3)OCI等于4,P/O为2671±123。(4)套袋实验表明紫花含笑能进行少量的自花传粉,人工异花授粉结实率和种子质量显著高于人工自花授粉和自然条件下,且不存在无融合生殖。(5)开花后36 h雌蕊群弯曲分泌粘液时,与含笑(M.figo)杂交结实率最高达85.7%,为最佳授粉时期。(6)有效访花者主要有东方蜜蜂、黑纹食蚜蝇和独角仙,访花频率易受天气影响。综上说明,紫花含笑繁育系统以异交为主,部分自交亲和需要传粉者,存在一定近交衰退,自然条件下结实率低,主要受传粉昆虫和雌雄异熟限制,最佳授粉期的确定能够提高其杂交成功率。该研究结果为紫花含笑资源保护和杂交育种奠定了基础。     

Pollination biology of the saprophytic speciesPetrosavia sakuraii (Makino) van Steenis in Central Japan

Pollination biology of the saprophytic speciesPetrosavia sakuraii was investigated in Central Japan. The flower opens at any time of the day, and the anthers of outer stamens dehisce after about one day and those of the inner stamens after two days. The stigmata on the semicarpous (nearly apocarpous) pistil, which are already receptive when the flower opens, are situated in the center of the flower and move gradually outwards to touch the anthers of outer stamens about five days after anthesis. The breeding experiments show that the stigmata are receptive at least for five days, and many seeds are produced through autonomous self-pollination. That means the nearly apocarpous pistil, which is in a primitive condition, is adapted to the autonomous self-pollination. The selfing rate including insect-mediated self-pollination is very high, and this sexual-reproductive system seems favorable to the saprophytic plant which is probably severely limited in its resources. However, cross-pollination also may be performed by the small bees of Lasioglossum and some other insects, even though they do not so frequently visit the flowers and the out-crossing rate is low.     

Reproductive strategies of diploid and polyploid populations of arcticDraba (Brassicaceae)

132 cultivated populations (2x–16x) of 15 arctic-alpine species ofDraba were investigated to clarify a possible relationship between reproductive strategies and polyploid evolution in the genus. The populations were exclusively sexual and produced viable seed after spontaneous self-pollination, but showed large variation both in traits promoting cross-pollination and in autogamous fruit and seed set. Traits promoting cross-pollination, e.g., floral display, protogyny, and delayed selfing, were positively correlated, and these traits were negatively correlated with autogamous fruit and seed set. All diploid and many polyploid populations had high autogamous seed set and small, unscented, non-protogynous, and rapidly selfing flowers. In contrast, all populations with low autogamous seed set and large, scented, and strongly protogynous flowers with distinctly delayed selfing were polyploid. These results are consistent with those previously obtained from enzyme electrophoresis, suggesting that the genetically depauperate diploids are extreme inbreeders and that the highly fixed-heterozygous polyploids vary from extreme inbreeders to mixed maters. The reproductive data lend additional support to the hypothesis that allopolyploidy in arcticDraba serves as an escape from genetic depauperation caused by uniparental inbreeding at the diploid level.     

Factors affecting the fruiting of bilberries: an analysis of categorical data set

The present study is an application of categorical data analysis in ecological research. The approach is based on logistic regression following an exploratory graphical analysis. The material was collected in an extensive forest inventory in which a set of observations was made by eye in a stratified random sample of 262 mature upland forest stands in South Finland. The problem was to interpret the variation in the fertility of the tillers of bilberry (Vaccinium myrtillus). In the field, the fertility was recorded as a three-class ordinal variable. The information available for the interpretation included the visually estimated density of the tree crowns, the soil fertility class determined using Cajander's forest site types and the percent cover of V. myrtillus.The GLM framework was employed in successive stages of the data analysis in order to find a model to fit the data. For this, the three-class ordinal response variable was reduced to two classes: stands characterized by (a) sterile and (b) fertile bilberry tillers. Successful prediction of the distribution of these two types of forest stands was achieved with a logistic-regression model by using canopy coverage and soil fertility classes as predictor variables. The generalized linear modelling framework is suitable for studying many ecological problems even when only rough categorical estimates of environmental scalars are available.     

THE CONSEQUENCES OF SELF-POLLINATION IN ASCLEPIAS EXALTATA,A SELF-INCOMPATIBLE MILKWEED

We investigated the effect of self-pollination on fruit- and seed-set in Asclepias exaltata, a self-incompatible milkweed. A total of 1,380 hand-pollinations were performed on 138 flowers in each of five treatments. In each treatment, two pollinia were placed in adjacent stigmatic chambers, which transmitted pollen tubes to the same ovary. Treatments 1 and 2 involved simultaneous placement of two self-pollinia (treatment 1) or two cross-pollinia (treatment 2) into adjacent stigmatic chambers. No fruits were produced after self-pollinations, but 23% of the cross-pollinated flowers matured fruits. The three remaining experimental pollination treatments were self-pollination preceding cross-pollination by 24 hr (treatment 3), simultaneous self- and cross-pollination (treatment 4), and cross-pollination preceding self-pollination by 24 hr (treatment 5). Compared with that in flowers receiving only cross-pollen, fruit-set in treatments 3–5 was reduced 81%, 49%, and 29%, respectively. Seed-set was also significantly reduced in flowers receiving self-pollen 24 hr in advance of the cross-pollen. Using genetic markers, we observed that only 0.6% of seeds resulted from self-pollination. Our data strongly suggest that self-pollination in milkweeds not only wastes pollen but also greatly reduces the number of ovules and ovaries that might otherwise mature fruits and seeds after cross-pollination.     

Regeneration strategies of dominant boreal forest dwarf shrubs in response to selective removal of understorey layers

Abstract. The deciduous Vaccinium myrtillus and the evergreen Vaccinium vitis‐idaea were subjected to five removal treatments of understorey layers: control, removal of the moss layer, removal of the field layer, removal of both moss and field layers and removal of moss, field and humus layers. A second factor, sowing, was included to investigate sexual reproduction after disturbance. Density of new ramets and seedlings and growth of annual shoots were studied for the first two growing seasons, whereas cover was measured for five growing seasons after disturbance treatment. Initially, vegetative production of new ramets and species cover increased rapidly in all disturbed plots, except for the most severe treatment, in which production of new ramets was virtually absent throughout the study. Full recovery following removal of the field layer only or both field and moss layers was reached after four years for V. myrtillus. V. vitis‐idaea recovered more quickly, after one year (removal of field layer only) and four years (removal of field and moss layers). The relative growth of V. myrtillus and V. vitis‐idaea increased in the latter treatment in terms of production of annual shoots and length of annual shoots, respectively. Seedling density increased after sowing in the most severe treatment. The results underscore the importance of vegetative growth for recovery of these species at moderate‐level disturbances. The high rate of sexual reproduction in the most severe treatment implies that strong mechanical disturbance is needed to enhance the establishment of new genotypes in these species.     

Characterization of a Variant ofAutographa californicaNuclear Polyhedrosis Virus with a Nonfunctional ORF 603

A new genotypic variant ofAutographa californicanuclear polyhedrosis virus (AcMNPV), the V8 variant, was originally identified by an additionalHindIII site in theHindIII–F fragment. Insect bioassays of this variant displayed a decreased time of mortality compared with the L1 variant of AcMNPV inSpodoptera frugiperdalarvae but not inTrichoplusia nilarvae. A 1.8-kb region containing the 3′ end of ORF 5,lef-2,ORF 603, and the 5′ end of the polyhedrin gene (polh) of both L1 and V8 was sequenced. V8 exhibited extensive sequence variation in the region between the 3′ end oflef-2and the 5′ end ofpolh; V8 had six amino acid substitutions in thelef-2gene product and a nonfunctional ORF 603. A site-specific frameshift mutation in ORF 603 of the L1 variant was constructed to determine the effect of ORF 603 inS. frugiperdalarvae. Truncation of ORF 603 was found to decrease the time of mortality inS. frugiperdalarvae. The insect-selective toxin gene,tox34, was inserted into the V8 variant by direct cloning. The efficacy of this recombinant as a biopesticide was equivalent to similar L1 recombinants.     

Early Inbreeding Depression and Pollen Competition inCalluna vulgaris(L.) Hull.

We investigated whether partial self-sterility inCalluna vulgarisresultsfrom abortion of selfed offspring owing to inbreeding depressionor a late-acting self-incompatibility mechanism, and whetherself-pollen interferes with normal functioning of cross-pollen.Self-pollination resulted in 75% less seed set than cross-pollination.Self-pollen tubes reached ovaries and penetrated ovules as oftenas those of cross-pollen. Following self-pollination, examinationof the size of undeveloped seeds showed that at least 70% resultedfrom ovule fertilization and arrest of development occurredat various stages. All self-pollinated plants produced seedsand self-fertility varied among plants. These results indicatethat the reduced seed set observed in self-pollination is morelikely the result of inbreeding depression rather than a late-actingself-incompatibility system. The fecundity component of inbreedingdepression was high (0.762). Seed set was reduced by an averageof 40% when self-pollen was mixed with cross-pollen, comparedto pure cross-pollination. Using genetic markers, we found about20% of seeds resulted from self-pollination in mixed-pollinatedfruits.C. vulgarisis likely to experience self-pollination innature and our data suggest this will reduce the number of ovulesthat might otherwise mature after cross-pollination.Copyright1999 Annals of Botany Company Calluna vulgaris(heather), self-pollination, pollen tube, ovule fertilization, early inbreeding depression, pollen interference.