Wind pollination in high-mountain populations of Hormathophylla spinosa (Cruciferae)

In this paper we study aspects of the breeding system of Hormathophylla spinosa (Crucifcrae) to identify the factors responsible for seed production in the absence of insect pollinators. The pollinator-exclusion experiments show that H. spinosa, under natural conditions, does not produce seed by apomixis or spontaneous autogamy. H. spinosa appears to be self-incompatible but slightly geitonogamous. Thus, this plant species needs pollen vectors for reproduction. The results of the wind-exclusion experiments performed during two different years in two populations of H. spinosa support the hypothesis that the wind acts as a pollen vector; flowers excluded from the wind had a lower fruit set and female fertility than flowers excluded from all pollinator insects (winged and wingless). This generalist pollination system (insects and wind) permits this species to colonize and sustain viable populations in high mountains, where this species is the only woody shrub living above 3000 m above sea level in the Sierra Nevada.     

Effect of ovule position within the pod on the probability of seed production in Bauhinia ungulata (Fabaceae)

BACKGROUND AND AIMS: It has been claimed that ovules linearly ordered within a fruit differ in their probabilities of reaching maturity. This was investigated by studying the effect the position of an ovule within the pod has on seed abortion and seed production in Bauhinia ungulata. METHODS: Fruits collected during the dry seasons of 1999, 2000 and 2001 were opened, and the number, position and status of each ovule within the fruit were recorded. A GLM model was used to assess the effects of population, tree identity and ovule position within the pod on ovule fertilization, seed abortion, seed damage and seed maturation in two populations of B. ungulata. KEY RESULTS: Nearly 30% of the ovules were not fertilized in 1999; this percentage dropped to 5% the following two years. Seed abortion (50%) and seed damage (15%) were the same every year during the study period. Only 15% of the initial ovules developed into mature seeds in 1999; this value increased to 35% in 2000 and 2001. However, seed survivorship was dependent on the position of the ovule within the pod; non-fertilized and early aborted ovules were found more often near the basal end of the ovary. The frequency of seed damage was not affected by position. Mature seeds were found mainly in the stylar half of fruits, where ovules are likely to be fertilized by fast pollen tubes. CONCLUSIONS: The pattern of seed production in B. ungulata is non-random but is dependent upon the position of the ovule within the pod. The results suggest that the seeds produced within a fruit might differ in their vigour.     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

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.     

Ovary,seed and fruit ofRutidea (Rubiaceae,Pavetteae)

The ovary ofRutidea is bicarpellate and incompletely bilocular (septum between locules not continuous). A solitary campylotropous ovule, ascending from a basal placenta, occurs in each locule. Based on their orientation and degree of curving, three ovule types are distinguished. As a consequence of the abortion of one ovule, the drupaceous fruits are one-seeded. The incomplete septum allows the spherical seed to fill out the entire interior of the fruit. The seeds are deeply ruminate (Spigelia type). They grow very fast, producing folds and undulations (ruminations) which invade and totally occupy the second locule, almost enveloping the aborted ovule. Comparisons with otherRubiaceae (especiallyPavetteae), show that hemianatropy and campylotropy occur more often in theRubiaceae than hitherto accepted. The study corroborates the close affinity betweenRutidea andNichallea.     

Temporal patterns of ovule allocation,fruit set,and seed predation inAnemonopsis macrophylla (Ranunculaceae)

Ovule deployment patterns were studied inAnemonopsis macrophylla Sieb. & Zucc. to develop hypotheses about major selective forces underlying different temporal strategies. Carpel number per flower decreased significantly with relative flowering date. The trend was present at individual, population, and among-population, levels. Both ovule number per carpel and ripe seed weight decreased with later flowering date. Such temporal allocation patterns may be adaptive in late-flowering species with extended flowering patterns, because successively later flowers run a higher risk of lower resource availability and premature death due to deteriorating climatic conditions.     

Triterpenoid saponins with antifeedant activities from stem bark of Catunaregam spinosa (Rubiaceae) against Plutella xylostella (Plutellidae)

Seven triterpenoid saponins, including four new compounds, catunarosides A–D (1–4), and three known compounds, swartziatrioside (5), aralia-saponin V (6), araliasaponin IV (7) were isolated from the stem bark of Catunaregam spinosa, a Chinese mangrove associate. Their structures were elucidated on the basis of their spectral data and hydrolysis experiments. The antifeedant activities of compounds 1–7 against Plutella xylostella were also evaluated.     

Variation of pollen and ovule parameters among different ploidy levels ofCorydalis (Fumariaceae)

The size and number of pollen grains and ovules are compared between 20 populations of different ploidy levels in two self-incompatible species ofCorydalis (Fumariaceae), to test the presence of ploidy-related variation in these reproductive characters. In both species, higher ploidy levels are associated with larger pollen grains, ovules and corolla, but the number of pollen grains and ovules are not different between ploidy levels. The investment per flower is consequently larger at higher ploidy levels, but the flower number per individual is lower, suggesting that the mode of partition of the investment for sexual reproduction varies between different ploidy levels. InC. orthoceras, sex allocation estimated by pollen:ovule ratios in number and volume is more female-biased in polyploids than in diploids. In spite of these variations, the characters studied can not be used as indicators of ploidy level due to the large overlaps between the ploidy levels.     

The co-occurrence of Artemia parthenogenetica and Branchinella spinosa (Branchiopoda: Anostraca) in a saline pond of south eastern Italy

The co-occurrence of Artemia parthenogenetica Bowen & Sterling, 1978 and Branchinella spinosa Milne-Edwards, 1840, was previously only supposed, on the basis of finding their cysts in the same lagoon, but active stages were never recorded together. In a saline lake in south eastern Italy, populations of the two species were observed for two years. A. parthenogenetica was typical of the early wet season (October–January), while B. spinosa, even if it was present before, developed in the late part of the wet season (February–May). Adult A. parthenogenetica were recorded only in the season 1999–2000. The seasonal distribution of A. parthenogenetica is probably controlled by abiotic conditions (i.e., photoperiod, temperature, conductivity, and rainfall); however, competition between specimens of similar body size (e.g., adults of A. parthenogenetica, and juveniles of B. spinosa) could explain the lower numbers of B. spinosa population in winter. The discovery of cysts of both species deep in the sediment of the lake suggests a long-lasting co-occurrence of these two Anostraca.     

Competition and the compensatory regulation of fruit and seed set in the perennial herb Epilobium dodonaei (Onagraceae)

I studied the effects of competition and soil fertilization on variation of seed yield components of the outcrossing perennial Epilobium dodonaei Vill. using a randomized complete block design. Fertilization as a main effect was not significant. Competition, however, reduced the number of flower buds per shoot (-29%), the number of ovules per fruit (-12%), and the number of ripe fruits per shoot (-51%). In addition, competition caused an increase in abortion of flower buds. Consequently the fruit/flower bud ratio decreased from 0.71 to 0.47. Average seed mass was not reduced significantly by competition. Some of the negative effects of competition on fruit production were mitigated by fertilization. However, competition considerably reduced the proportion of late-aborted seeds, which resulted in an increase of the seed/ovule ratio from 0.31 to 0.49 (+58%). As a consequence of this compensation, the reduction in the number of seeds per shoot due to competition was not significant. At the level of the genet, competition had a strong effect on seed yield due to decreases in the number of shoots produced. I discuss patterns of regulation at consecutive levels of reproduction. Shoots of plants suffering from competition initially invested less in reproductive structures than the control and showed a higher abortion rate of less costly structures early in reproductive development, but also had a lower abortion rate later in their development. The observed compensatory pattern in yield components illustrates the fine-tuning regulation capacity at different levels of reproductive development in plants.     

Isozyme and seed protein phylogeny of the genusCitrullus (Cucurbitaceae)

Electrophoretic analysis of 26 enzyme coding genes was conducted on accessions of threeCitrullus species and the relatedPraecitrullus fistulosus andAcanthosicyos naudinianus. The isozyme phylogeny of the genusCitrullus and the related species was constructed based on pairwise measurements of the respective genetic distances between the species and races.P. fistulosus andA. naudinianus form two distinct outgroups toCitrullus which is characterized by two main clusters: The first includes twoC. colocynthis races and the second,C. lanatus andC. lanatus var.citroides, which are more closely related to each other than they are toC. ecirrhosus. The isozyme phylogeny is consistent with the variability in six seed protein bands and with the crossability relations among the examined species.     

Selective fruit and seed maturation in Asphodelus albus Miller (Liliaceae)

The fruiting patterns of the rhizomatous perennial Asphodelus albus Miller (Liliaceae) were studied in five populations during 1989 and in one population over 3 years (1988–1990). Fruit/flower (Fr/Fl) ratio and seed/ovule (S/O) ratio varied markedly between populations. Although there were differences between years within the population studied over 3 years, these variations, although statistically signifcant, were less important than those between populations. Neither flowering phenology nor plant size influenced Fr/Fl or S/O ratios. Field experiments tested whether fruit and seed set were pollenor resource-limited. Hand-pollination had no effect on Fr/Fl or S/O ratios, but the reduction of resources by defoliation at the time of flowering decreased both relative components of fecundity. Manipulation of resources by hand-thinning flowers and tiny fruits had no demonstrable effect on these ratios, although brood size of individual fruits was significantly affected. It may be concluded that fruit maturation is resource-limited rather than pollen-limited. Most of the fruits aborted early in the fruiting season, and fruits with higher numbers of developing seeds had a lower probability of abortion than fruits with fewer seeds. Analyses of position effects revealed that the fruits in lower positions in the inflorescence matured preferentially. Furthermore, the two ovules in the same carpel tended either both to fail or both to develop into seeds. The plant's ability to selectively mature only high quality embryos may be limited.     

In vitro propagation of caper (Capparis spinosa L.)

Summary A twenty fold multiplication per twenty days of caper was achieved by culturing nodal shoot segments in the presence of BAP (4 μM) plus IAA (0.3 μM) and GA₃ (0.3 μM). The use of a modified MS medium facilitated this response. Plantlet regeneration was induced on single shoots taken from proliferating clusters subcultured for 20 days on a reduced BAP (2 μM) without auxin and gibberellin Higher rooting responses (70%) were obtained after a 20-day incubation period in darkness on solid half-strength MS1 medium plus IAA (30 μM), followed by a subsequent 20 day culture period on half-strength MSI basal medium. Proliferation was mainly due to axillary shoot-bud development as revealed by histological studies. The extensive meristematic activities observed indicated the enormous morphological potential of this species.     

Germination behaviour in populations ofCapsella bursa-pastoris (Cruciferae)

Germination behaviour of variousCapsella bursa-pastoris populations collected from Scandinavia, Middle Europe and the Alps, was tested in unheated, non-illuminated greenhouses (46 populations) and in growth chambers using 5–7 alternating temperature regimes (16 populations). For all populations, the influence of temperature on germination rate is straightforward: the higher the temperature, the greater the germination. Germination capacity, however, may depend on the geographical region. There is also a strong seed age effect on both, rate and capacity of germination. Once dormancy was broken, seeds from all populations were able to germinate over the entire range of temperatures. Some populations revealed a more or less pronounced temperature optimum for germination capacity, others germinated equally well over the entire temperature range. This indicates genetic heterogeneity between populations. However, no correlation between germinability and any environmental pattern was detected. The data indicate thatCapsella bursa-pastoris has adopted a germination strategy which includes a broad temperature tolerance. Germination of wildCapsella plants seems to be regulated by the factors contributing to the inception and breaking of dormancy which depend on pre- and postharvest conditions. Adaptation in germination behaviour inCapsella bursa-pastoris is different from that in other life history traits (flowering behaviour, growth form parameters).     

Multivariate and cladistic analyses of the purple-flowered species ofErysimum (Cruciferae) from the Iberian Peninsula

A taxonomically difficult purple-flowered group within the genusErysimum, restricted to the Iberian Peninsula, is analyzed by multivariate and cladistic analyses. 51 specimens have been scored for 14 characters. Both principal components and discriminant analyses provide support to the recognition of the five species considered by the author, namelyE. linifolium, E. lagascae, E. baeticum, E. popovii, andE. cazorlense. Cladistic analysis, using 7 characters resulted in a single most parsimonious cladogram containing no homoplasies. The pattern of morphologic divergence follows a clear NW.-SE. trend, which is congruent with the topology of the cladogram. This trend significantly affects growth-form as well as fruit characters, both providing the main grounds for species recognition. The different behavior and significance of several characters in both kinds of analysis is discussed. The co-occurrence of morphologically similar individuals differing in the flower color is discussed, too. Possible explanations for this phenomenon involve hybridization in a wide sense or, alternatively, rejecting the assumption of monophyly for the group.     

The Association between Medusa Lychnorhiza lucerna (Scyphomedusae, Rhizostomeae) and Decapod Libinia spinosa (Brachyura, Majidae) Recorded for the First Time in Neritic Waters of Argentina

The paper discusses the first record of the association between the medusa Lychnoriza lucerna Haeckel, 1880 and the decapod Libinia spinosa Milne-Edwards, 1934 in neritic waters of Argentina. Our results allow us to conclude that the symbiotic relation between these two species is a common phenomenon. The crab L. spinosa is located in the subgenital pockets of L. lucerna such that one-half of its cephalothorax is exposed to the outside. The crabs feed at the expense of the medusa, ingesting some portion of its food. A few specimens of the crab have been found, mostly females; the only male was of a somewhat larger size.     

Factors affecting in vitro microrhizome production in turmeric

In vitro microrhizome production was obtained in turmeric (Curcuma longa Linn.). Freshly sprouted buds with small rhizome portions excised from stored mature rhizomes were cultured on semi-solid culture initiation medium –- MS basal medium + 0.88 M BAP (6-benzylaminopurine) + 0.92 M kinetin + 5% coconut water + 2% sucrose + 0.5% agar –- resulting in bud elongation. Multiple shoots were produced from these elongated buds by culturing in liquid shoot multiplication medium –- MS basal medium + 2.2 M BAP + 0.92 M kinetin + 5% coconut water + 2% sucrose –- at 25±1°C and 16-h light (at 11.7 mol m^–2 s^–1)/8-h dark cycles. Clumps of four to five multiple shoots/single shoots were used in various experiments. Cultures were incubated in the dark at 25±1°C. Half strength MS basal medium supplemented with 80 g l^–1 sucrose was found to be optimal for microrhizome production. Cytokinin BAP had an inhibitory effect on microrhizome production. At the highest concentration of BAP tried (35.2 M) microrhizome production was totally inhibited. Microrhizome production depended on the size of the multiple shoots used. Microrhizomes produced were of a wide range in size (0.1–2.0 g) and, readily regenerated when isolated and cultured in vitro on culture initiation medium or shoot multiplication medium. Under in vivo conditions, small (0.1–0.4 g), medium (0.41–0.8 g) and big (>0.81 g) microrhizomes regenerated. Plantlets developed from big microrhizomes grew faster.     

Factors affecting progeny production of Anaphes iole

Anaphes iole Girault(Hymenoptera: Mymaridae) is a native, solitaryegg parasitoid of Lygus spp.(Heteroptera: Miridae) in North America. Current research is considering factors thatmay optimize the in-vivo rearing of A.iole using Lygus hesperus Knight(Heteroptera: Miridae) as host. The effects ofhost density, day of oviposition and foodpresence, parasitoid age and mate presence onproduction of A. iole progeny weredetermined in this study. After exposingindividual parasitoids to host patches for 24h, the percentage of hosts containing a latestage A. iole pupa wassignificantly greater at a moderate hostdensity (41–70 eggs per patch) than at a lowdensity (10–40 eggs). But, no differenceswere detected between moderate and high density(71–100 eggs) or between high and low densitytreatments. More adult progeny were generatedby females (of variable age; 0–2 d old) onthe first day of oviposition rather than thesecond day, regardless of food presence. Progeny sex ratio was decidedly male-biased onthe second day. Female age (0 d vs 1 d old)had a marginal effect on production in 24 h; 0d old females tended to generate more adultprogeny than 1 d old females. Overall, thisresearch suggests that exposing newly-emerged,unfed, mated A. iole females to amoderate to high host density for 1 to 2 dcould lead to time-efficient production ofadult progeny in an in-vivo rearing system.     

Non-additive effects of herbivores and pollinators on Erysimum mediohispanicum (Cruciferae) fitness

In this study, the non-additivity of effects of herbivores and pollinator on fitness of the plant Erysimum mediohispanicum (Cruciferae) has been experimentally tested. The abundance and diversity of the pollinator assemblage of plants excluded from and exposed to mammalian herbivores, and the combined effect of pollinators and herbivores on plant reproduction were determined over a period of 2 years. Pollinator abundance was higher and diversity was lower on plants excluded from herbivores. Furthermore, the experimental exclusions demonstrated that both pollinators and herbivores affected plant fitness, but their effects were not independent. Herbivores only had a detrimental effect on plant fitness when pollinators were present. Similarly, pollinators enhanced fitness only when herbivores were excluded. This outcome demonstrates that the importance of pollinators for plant fitness depends on the occurrence of herbivores, and suggests that herbivores may hamper pollinator-mediated adaptation in plants.