Synflorescences of species related to Schizachyrium condensatum (Poaceae)

The synflorescences of species related to Schizachyrium condensatum are typologically characterized. This species presents floriferous shoots with complex systems of ramification. In all species the inflorescence is polytelic and truncate. The typological pattern has been described and presents differences in the following parameters: length and shape of the synflorescence, paracladia of the trophotagma subzone and short paracladia subzone. A comparative analysis of the variations observed in the structure of the synflorescence is included.     

Modular growth and functional heterophylly of the phreatophyte Ziziphus lotus: A trait-based study

The variation of plant functional traits, from the cell to the whole-plant level, is a central question in trait-based ecology with regard to understanding ecological strategies and adaptations that result from environmental drivers. Here, we analyzed whole-plant and leaf traits of the phreatophyte Ziziphus lotus (L.) Lam., a long-lived shrub that dominates one of the few terrestrial groundwater-dependent ecosystems (GDEs) in Mediterranean Basin drylands. We (a) assessed architectural traits and growth patterns, (b) analyzed leaf morpho-functional traits (specific leaf area [SLA] and stomata pore index [SPI]) and physiological traits (gas exchange rates), as well as their variations within individuals, and (c) evaluated temporal variations in modular growth (i.e., sequential iteration of structural units) between growing seasons and in leaf traits within seasons. Z. lotus' growth pattern was based on the repetition of modules composed of shoots (short and long) and branches (flowering and plagiotropic) that promoted a functional differentiation between vegetative and reproductive structures, respectively. We identified morpho-functionally distinct leaves (i.e., heterophylly) borne on different types of branches. Leaves on flowering branches had higher SLA and water use efficiency (WUEi), but lower SPI and transpiration rates than leaves on vegetative ones. We also observed trade-offs in the elongation of vegetative and flowering structures between growing seasons: the shorter the long shoots, the larger the flowering branches. The modular differentiation and heterophylly of Z. lotus might contribute to prioritizing the investment of resources of this phreatophyte, either for growth or reproduction, and could improve the efficiency in uptake and conservation of resources in drylands.     

Cost and probability of flowering at the shoot level in relation to variability in shoot size within the crown of Vaccinium hirtum (Ericaceae)

Size-related variation in the cost and probability of flowering among shoots within a crown of Vaccinium hirtum was investigated to clarify patterns and regulation of flowering at the shoot-module level, below the level of the individual. The apices of previous-year shoots differentiated into current-year shoots vegetatively (vegetative branches) or became reproductive by developing inflorescences (reproductive branches). Length growth and fate of current-year shoots were determined, and the future potential for reproduction was estimated using a matrix model of shoot dynamics. Reproductive branches had fewer current-year shoots and shorter total shoot lengths and thus had a reduced potential for reproduction compared with vegetative branches, indicating the cost of flowering at the shoot level. This cost of flowering was higher in longer shoots. The probability of the initiation of flowering in a shoot increased with increasing shoot length in shorter shoots, reached a maximum in medium-sized shoots, and decreased in longer shoots. The size-related changes in the probability of flowering at the shoot level can be largely explained by the size-dependent changes in shoot-level resource availability and cost of flowering.     

Comparative morphology of acrogenous branch systems and phylogenetic considerations. II. Angiosperms

A concept for a primitive angiospermous branch system is given in order to have a starting point for the derivation of the diverse and highly differentiated branch systems observed in contemporary angiosperms. Hitherto Troll's (1964, 1969) comparative study of the synflorescences in this plant group — developed out predominantly on herbaceous plants — was the most comprehensive and sophisticated treatment dealing with branch systems. Unfortunately, the work on tropical tree architecture by Hallé et al. (1978) has no reference to the classical studies of Troll and his pupils. Thus Müller-Doblies and Weberling (1984) emphasized the high degree of terminological incompatibility between the two works. Angiosperms are seen as a monophyletic plant division. Consequently, the branch system of the first primitive angiosperms must be the starting point in the evolution of the abundant diversity of branch systems and growth forms of modern angiosperms. If it is accepted that primitive angiospermous shoots were terminated by a large flower, one may assume, that the reproductive end of the shoot was enriched by paracladia early in evolution, thereby developing a terminal inflorescence instead of a single flower. Thus the primitive shoot unit was divided into a basal vegetative region — the trophotagma, branching retardively — and the reproductive terminal region, the anthotagma, branching simultaneously. It is demonstrated through a selection of different examples, that the construction of such a system possesses the options for several modifications, enabling the evolution of the abundant diversity of branch systems which characterizes contemporary angiosperms.     

Branch geometry in Cornus kousa (Cornaceae): computer simulations

Computer simulations similar to actual trees were constructed using simple branching rules. Branch orientation with respect to the direction of gravity was a fundamental consideration. In Cornus kousa BUERG. ex HANCE, several types of branches develop from winter buds, varying from orthotropic shoots to plagiotropic ones. Based on actual observations and measurements of branching structures with a wide range of orientations, we made a flexible geometrical model consisting of five forking branches that varied in outgrowth depending on the direction of the shoot with respect to gravity. Repetition of the branching by computer generated a realistic tree pattern, which was close to the shape of a young C. kousa tree. Reproductive shoots seem to be under a branching rule that was a modification of vegetative branching, although the reproductive branch size was considerably smaller than the vegetative one, and reproductive branching was bifurcated instead of five-forked. We conclude that all branchings in orthotropic and plagiotropic shoots in the vegetative phase and shoots in the reproductive phase are formed under the same branching rule, but each has different parameter values.     

Inflorescence structure in species of Spartina Schreb. (Poaceae: Chloridoideae: Cynodonteae)

The typology developed by Troll was followed to describe inflorescence structures for 15 species of Spartina. In all species here studied, truncation of the terminal spikelet of the main axis and primary paracladium was observed. The truncation can also involve the short paracladia subzone so that the inflorescence is confined to just the long paracladia subzone. A great homogenization of paracladia and maximum ramification degree limited generally to the second order of branching are distinctive characteristics of the genus. Proximal paracladia with third-order branching were found in only three specimens, and in these exceptional cases, the homogenization is partial. Sometimes, in some species, a subzone of long and short paracladia can be distinguished. The absence of trophotagma paracladia in all the species studied was verified. The variation in the structure of the inflorescence among species is due to the differences in the number of short paracladia, long paracladia, total number of primary paracladia and also in the angle of divergence of the long paracladia from the main axis. The latter, in addition to variations in the intercalary growth of the internodes produces modifications in the general appearance of the inflorescences. The systematic and taxonomic value of the inflorescences in Spartina is discussed.     

How endangered is sexual reproduction of high-mountain plants by summer frosts? Frost resistance, frequency of frost events and risk assessment

In temperate-zone mountains, summer frosts usually occur during unpredictable cold spells with snow-falls. Earlier studies have shown that vegetative aboveground organs of most high-mountain plants tolerate extracellular ice in the active state. However, little is known about the impact of frost on reproductive development and reproductive success. In common plant species from the European Alps (Cerastium uniflorum, Loiseleuria procumbens, Ranunculus glacialis, Rhododendron ferrugineum, Saxifraga bryoides, S. moschata, S. caesia), differing in growth form, altitudinal distribution and phenology, frost resistance of reproductive and vegetative shoots was assessed in different reproductive stages. Intact plants were exposed to simulated night frosts between ?2 and ?14 °C in temperature-controlled freezers. Nucleation temperatures, freezing damage and subsequent reproductive success (fruit and seed set, seed germination) were determined. During all reproductive stages, reproductive shoots were significantly less frost resistant than vegetative shoots (mean difference for LT₅₀ ?4.2 ± 2.7 K). In most species, reproductive shoots were ice tolerant before bolting and during fruiting (mean LT₅₀ ?7 and ?5.7 °C), but were ice sensitive during bolting and anthesis (mean LT₅₀ around ?4 °C). Only R. glacialis remained ice tolerant during all reproductive stages. Frost injury in reproductive shoots usually led to full fruit loss. Reproductive success of frost-treated but undamaged shoots did not differ significantly from control values. Assessing the frost damage risk on the basis of summer frost frequency and frost resistance shows that, in the alpine zone, low-statured species are rarely endangered as long as they are protected by snow. The situation is different in the subnival and nival zone, where frost-sensitive reproductive shoots may become frost damaged even when covered by snow. Unprotected individuals are at high risk of suffering from frost damage, particularly at higher elevations. It appears that ice tolerance in reproductive structures is an advantage but not an absolute precondition for colonizing high altitudes with frequent frost events.     

Trade-offs between vegetative growth and acorn production in Quercus lobata during a mast year: the relevance of crop size and hierarchical level within the canopy

The concept of trade-offs between reproduction and other fitness traits is a fundamental principle of life history theory. For many plant species, the cost of sexual reproduction affects vegetative growth in years of high seed production through the allocation of resources to reproduction at different hierarchical levels of canopy organization. We have examined these tradeoffs at the shoot and branch level in an endemic California oak, Quercus lobata, during a mast year. To determine whether acorn production caused a reduction in vegetative growth, we studied trees that were high and low acorn producers, respectively. We observed that in both low and high acorn producers, shoots without acorns located adjacent to reproductive shoots showed reduced vegetative growth but that reduced branch-level growth on acorn-bearing branches occurred only in low acorn producers. The availability of local resources, measured as previous year growth, was the main factor determining acorn biomass. These findings show that the costs of reproduction varied among hierarchical levels, suggesting some degree of physiological autonomy of shoots in terms of acorn production. Costs also differed among trees with different acorn crops, suggesting that trees with large acorn crops had more available resources to allocate for growth and acorn production and to compensate for immediate local costs of seed production. These findings provide new insight into the proximate mechanisms for mast-seeding as a reproductive strategy.     

Probleme der Infloreszenztypologie vonW. Troll

In contrast toW. Troll's typology of inflorescences which aims at more or less rigid, well defined types, this investigation accentuates the processes that constitute the evolutionary transformations leading from one typical form to another.Troll divided the inflorescences into the two types of monotelic and polytelic synflorescences, the first with a terminal flower on the main axis, the latter with a homogeneous florescence on the indeterminate axis. Both forms are enriched by proximal branches which repeat the structure of the main axis (paracladia). The evolutionary processes leading from the more primitive monotelic type to the advanced polytelic type are truncation (loss of the terminal flower) and homogenization of the distal branches, which thus form a homogeneous florescence. A closer survey of the polytelic groups reveals the fact that, usingTroll's criteria, the same distinction can be found within these groups themselves. Loss of the terminal florescence (truncation of 2nd and higher degree) as well as homogenization of the distal paracladia may lead to florescence-like units of higher complexity. Examples can be found inAsteraceae (Figs. 1 and 2),Fabaceae (Fig. 3 a),Mimosaceae (Fig. 3 b),Acanthaceae, and also in Monocots, as exemplified by theMarantaceae (Figs. 4 and 5). The so-called racemization (inversion of efflorescences from basipetalous to acropetalous) may be mentioned as a third element of transformation, emphasizing the unity of the florescences.—In consequence, there are more organizational levels than reflected in the twoTroll types. The polytelic type comprises several degrees of truncation and homogenization, the basis for a reasonable organizational analysis should therefore be the degree of ramification of flowering branches rather than the mere question of a terminal flower on the main shoot axis (Fig. 6). On the other hand the three processes of truncation, homogenization and racemization are evolutionary transformations that may occur independently from one another, thus giving rise to a large number of variations, which can not be satisfactorily interpreted by exactly defined types. On the basis of these considerations the question of homologous parts in inflorescences is reviewed. The homology of partial florescences and paracladia is accentuated contrary toTroll's interpretation (Fig. 7). Homogenization as an evolutionary trend may transform paracladia of different degree of ramification, leading to one-flowered units on the one side and to highly complex structures as in theMarantaceae on the other.

     

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae)

To elucidate the significance of the simultaneous growth of vegetative and reproductive organs in the prostrate annual Chamaesyce maculata (L.) Small (Euphorbiaceae) from the standpoint of meristem allocation, we investigated plant architecture, meristem allocation, and the spatial and temporal patterns in vegetative growth and reproduction in the reproductive stage. The numbers of secondary and tertiary shoots successively increased by branching in the reproductive stage, and the sum of shoot length was greater in secondary shoots than in primary shoots. The specific shoot length (shoot length per shoot biomass) was greater in lateral shoots than in primary shoots, indicating efficient lateral shoot elongation. The internode length was shorter in secondary shoots than in primary shoots, increasing the number of nodes per shoot length in secondary shoots. Many nodes on a shoot generated two meristems, one of which committed to a flower and one to a lateral shoot. The number of reproductive meristems was greatest in tertiary shoots, and 96% of total reproductive meristems on shoots were generated in lateral shoots. On almost all nodes, the reproductive meristem developed into a flower, and 95–98% of the flowers produced a fruit. Therefore, vegetative growth by branching in the reproductive stage contributed to the increase in reproductive outputs. From the standpoint of meristem allocation, the simultaneous growth of vegetative and reproductive organs in prostrate plant species might be important for increasing the number of growth and reproductive meristems, resulting in the increase in reproductive outputs.     

Refining the biological factors affecting virulence of Botryosphaeriaceae on grapevines

Botryosphaeriaceae isolates of six species were assessed for their potential to infect grapevine tissues other than their tissues of isolation, primarily to determine sources of inocula that could contribute to bunch rot. Pathogenicity tests were conducted in vitro on berries and wood and in vivo on dormant buds of cultivars Chardonnay and Shiraz in glasshouse and field experiments. Tissue specificity and variation in virulence for different isolates was assessed. All isolates were able to infect and cause symptoms on detached 1‐year‐old canes and mature berries. Virulence was not affected by origin tissue and varied between isolates and within species. Inoculation of dormant buds did not affect bud burst or further development of shoots and fruit, however, a small number of Botryosphaeriaceae were reisolated from bunches at later growth stages. We conclude that Botryosphaeriaceae species are important pathogens of both the vegetative tissues and wood of grapevines. Grapevine wood infected with Botryosphaeriaceae could act as a source of inoculum for reproductive and vegetative tissue. Equally, the vegetative and reproductive tissues can also act as inoculum sources for wood infection. Therefore, all sources of inocula should be taken into consideration when developing management strategies for Botryosphaeria bunch rot and Botryosphaeria canker diseases.     

腾格里沙漠人工固沙植被中油蒿的生长及生物量分配动态

植物的资源分配模式反映了对环境的生态适应对策。2007年整个生长季, 采用生物量法对腾格里沙漠东南缘固沙植被区半灌木油蒿(Artemisia ordosica)地上部分各器官的生长及资源分配格局动态进行了研究。结果表明: 不同时期各器官的生长速率不同, 光合产物在各器官中的分配也不是等量的, 而是按一定的顺序在不同时期有不同的分配中心; 2007年油蒿的营养生长、繁殖输出、生殖枝大小都显著大于年降水量不足其一半的年份, 而繁殖分配和头状花序大小没有差异; 营养器官生物量大的油蒿总的繁殖输出也大, 但生殖期内营养生长和生殖生长既不同时也不等速, 表明资源分配的权衡(Trade-off)是存在的; 固沙植被建立以后, 随着时间延长, 油蒿的当年总生物量、繁殖输出、繁殖器官生物量分配有减小的趋势, 但不显著。     

Sex allocation and reproductive success in the andromonoecious perennial Solanum carolinense (Solanaceae). I. Female

Relative allocation of resources to growth vs. reproduction has long been known to be an important determinant of reproductive success. The importance of variation in allocation to different structures within reproductive allocation is somewhat less clear. This study was designed to elucidate the importance of allocation to vegetative vs. reproductive functions, and allocation within reproductive functions (sex allocation), to realized female success in an andromonoecious plant, Solanum carolinense. Allocation measurements were taken on plants in experimental arrays exposed to natural pollination conditions. These measurements included total flower number, the proportion of flowers that were male, flower size, and vegetative size. Flower number explained the majority of the variation among individuals in their success-that is, there was strong selection for increased flower production. There was also selection to decrease the proportion of flowers that were male, but neither flower size nor vegetative size (a measure of overall resource availability) were direct determinants of female success. After Bonferroni corrections for multiple comparisons, most phenotypic correlations among the traits measured were nonsignificant. Thus, in this andromonoecious species there is not a strong relationship between resource availability (vegetative size) and female success, and female success is instead determined by the relative production of the two different flower types.     

Patterns of vegetative growth and reproduction in relation to branch orders: the plant as a spatially structured population

The patterns of vegetative growth and reproduction in relation to orders of terminal branches were examined in the evergreen woody plant, Eurya japonica. The branch order number was determined centrifugally. The trunk was given order number 1; branches issuing directly from the trunk were order 2; branches growing on order 2 branches were order 3, and so on. The results of this study show the differential patterns of vegetative growth and reproduction in relation to the branch orders. Lower-order shoots of terminal branches grew more, but produced few flowers. On the other hand, for the higher-order terminal branches, shoot growth was very limited but flowering was more intense. The results show that a tree can be interpreted not as a mere population of equivalent modules but as a spatially structured population. Thus, it is essential to consider the position of modules within the branch system when patterns of vegetative growth and reproduction are examined. It is hypothesized that the difference in the opportunity cost in relation to the branch orders is the main cause of the spatial structure for patterns of vegetative growth and reproduction. Furthermore, for same-order terminal branches, current-year shoot elongation was independent of flowering intensity. These results suggest that this species only invests resources in reproduction that are surplus to its requirements for the functions associated with survival, such as growth and/or storage. Received: 22 July 1999 / Accepted: 12 January 2000     

Within-tree and among-tree variation in growth of Epirrita autumnata on mountain birch leaves

Abstract.

• 1 We studied among-tree and within-tree variation in the growth of larvae of a geometrid, Epirrita autumnata, on mountain birch (Betula pubescens ssp. tortuosa) leaves at different hierarchical levels: among trees (genets), among ramets within trees, among branches within ramets, among shoots within branches and among leaves within shoots. We used only short shoot leaves, which burst simultaneously in spring.
• 2 Trees accounted for most of the variation in larval growth rate, but there was substantial variation also among ramets within trees, among branches within ramets, and among short shoots within branches. Variation among leaves within short shoots was negligible. When the probabilities from different experiments were combined, the differences were statistically significant at the tree, ramet and branch levels, and approached significance at the shoot level.
• 3 In different experiments, larval growth was from 9% to 54% lower on the worst tree than on the best tree of the experiment. On average, larval growth rate was in different experiments from 11% to 32% lower on the worst ramet than on the best ramet within tree, from 8% to 18% lower on the worst than on the best branch within ramet, and from 12% to 30% lower on the worst than on the best shoot within branch.
• 4 The amount of among-and within-tree variation shown by our results may have ecological and evolutionary implications: among-tree variation should select for discrimination by ovipositing females and dispersing larvae, within-tree variation should select also for optimal foraging behaviour of larvae.

     

Costs of reproduction as related to the timing of phenological phases in the dioecious Shrub Pistacia lentiscus L

Females of woody dioecious species usually devote more resources to reproduction than males. This may lead to a decrease in female survival and growth. The costs of reproduction, however, can be lightened through a number of mechanisms, as for example avoiding the temporal coincidence of reproduction and vegetative growth. The aim of this study was to evaluate whether males and females of P. lentiscus differ in the timing of their vegetative growth, and to assess whether the sequencing of vegetative growth and reproduction reduces reproductive costs. We monitored phenology in males and females. We also compared male and female allocation of nutrients and biomass in the branch, and the developmental stability of the growing shoots. We did this both prior to and at the end of the fruiting period. Males and females showed similar vegetative and flowering phenologies. Males invested more biomass in flowering, but the sexes showed equal vegetative biomass and nutrient content prior to the fruiting period. In female branches, no trade-off was found between fruit load and current-year vegetative growth. In P. lentiscus, avoiding the overlap of flowering, vegetative growth and fruiting probably contributes to reduce the immediate costs of reproductive efforts, both in males and females.     

Timing of reproductive and vegetative development in Saxifraga oppositifolia in an alpine and a subnival climate

Morphogenesis of floral structures, dynamics of reproductive development from floral initiation until fruit maturation, and leaf turnover in vegetative short-stem shoots of Saxifraga oppositifolia were studied in three consecutive years at an alpine site (2300 m) and at an early- and late-thawing subnival site (2650 m) in the Austrian Alps. Marked differences in the timing and progression of reproductive and vegetative development occurred: individuals of the alpine population required a four-month growing season to complete reproductive development and initiate new flower buds, whereas later thawing individuals from the subnival sites attained the same structural and functional state within only two and a half months. Reproductive and vegetative development were not strictly correlated because timing of flowering, seed development, and shoot growth depended mainly on the date of snowmelt, whereas the initiation of flower primordia was evidently controlled by photoperiod. Floral induction occurred during June and July, from which a critical day length for primary floral induction of about 15 h could be inferred. Preformed flower buds overwinter in a pre-meiotic state and meiosis starts immediately after snowmelt in spring. Vegetative short-stem shoots performed a full leaf turnover within a growing season: 16 (+/-0.8 SE) new leaves per shoot developed in alpine and early-thawing subnival individuals and 12 (+/-1.2 SE) leaves in late-thawing subnival individuals. New leaf primordia emerged continuously from snowmelt until late autumn, even when plants were temporarily covered with snow. Differences in the developmental dynamics between the alpine and subnival population were independent of site temperatures, and are probably the result of ecotypic adaptation to differences in growing season length.     

Distribution of plant‐dwelling spiders: Inflorescences versus vegetative branches

Abstract The influence of the architecture of vegetative branches on the distribution of plant‐dwelling spiders has been intensively studied, and the effects on the aggregation of individuals in several spider species on plants include variation in prey abundance, availability of predator‐free refuges and smoother microclimate conditions. The emergence of inflorescences at the reproductive time of the plants changes branch architecture, and could provide higher prey abundance for the spiders. The distribution of spiders between inflorescences and vegetative branches was compared on four widespread plant species in a Brazilian savannah‐like system. Inflorescences attracted more spiders than vegetative branches for all plant species sampled. The influence of branch type (inflorescence and vegetative) on spider distribution was also evaluated by monitoring branches of Baccharis dracunculifolia DC. in vegetative and flowering periods for 1 year, and through a field experiment carried out during the same period where artificial inflorescences were available for spider colonization. Artificial inflorescences attached to B. dracunculifolia branches attracted more spiders than non‐manipulated vegetative branches for most of the year. However, this pattern differed among spider guilds. Foliage‐runners and stalkers occurred preferentially on artificial inflorescences relative to control branches. The frequencies of ambushers and web‐builders were not significantly different between treatment and control branches. However, most ambush spiders (65%) occurred only during the flowering period of B. dracunculifolia, suggesting that this guild was influenced only by natural inflorescences. The experimental treatment also influenced the size distribution of spiders: larger spiders were more abundant on artificial inflorescences than on vegetative branches. The hypothesis that habitat architecture can influence the spider assemblage was supported. In addition, our observational and experimental data strongly suggest that inflorescences can be a higher quality microhabitat than non‐reproductive branches for most plant‐dwelling spiders.