ULTRASTRUCTURE OF DEVELOPING AND MATURE NONARTICULATED LATICIFERS IN THE MILKWEED ASCLEPIAS SYRIACA L. (ASCLEPIADACEAE)

The ultrastructure of developing and mature nonarticulated laticifers in Asclepias syriaca L. (the common milkweed) was studied by conventional fixation and staining techniques and by osmium impregnation techniques. The mature laticifer protoplast in A. syriaca possesses a large central vacuole with an intact vacuolar membrane. Formation of this vacuole apparently results from dilation and subsequent enlargement of endoplasmic reticulum and possibly in part by fusion of smaller vacuoles and limited cellular-lytic autophagy. Widespread digestion or autophagy of cytoplasm within vacuoles is not evident. Nuclei, mitochondria, dictyosomes, and small vesicles are the most prominent components distributed in the peripheral cytoplasm. Plastids appear to degenerate as the laticifer matures. The specialized cellular component, latex, which is the vacuolar content of the laticifer, is interpreted to be produced in the cytoplasm and subsequently incorporated into the large central vacuole. Rubber globules, the most prominent latex component, are surrounded by a membrane that does not have a trilaminate structure. Globules are associated with an electron-dense fibrillar component in the vacuole.     

ENVIRONMENTAL INFLUENCES ON NECTAR PRODUCTION IN MILKWEEDS (ASCLEPIAS SYRIACA AND A. EXALTATA)

Cross-comparisons of nectar production data are complicated because different workers use bags made of various materials to exclude animal visitors. Using clonal populations of Asclepias syriaca and A. exaltata in northern Virginia, we carefully measured the effects of four bagging treatments (bridal veil, pellon, paper, plastic) on microenvironment (temperature, relative humidity) and nectar production (volume, concentration, sucrose amount) over the course of a day. In general, bridal veil bags changed the microenvironment least relative to unbagged controls. Plastic bags resulted in higher temperatures and constantly higher relative humidities. Temperature and relative humidity were also elevated, though less dramatically, in paper and pellon bags. Under more humid conditions, flowers contained larger volumes of more dilute nectar. Therefore, researchers who wish to obtain nectar production data that reflect natural field conditions should use bridal veil, or a material with similar properties, to bag inflorescences. We also performed a watering experiment, involving the addition of the equivalent of a 10-cm rain to the A. syriaca plot. After watering, nectar volumes and sucrose amounts were increased approximately twofold.     

BIFURCATION OF THE STEM APEX IN ASCLEPIAS SYRIACA

In Asclepias syriaca the overall inflorescence is an anthoclad in which the peduncles are non-axillary, each occurring about 60° away from the axil of a leaf. Ontogenetically, a peduncle is initiated when the stem apex expands laterally and bifurcates into separate apices, neither of which is subtended by any type of organ. One of the two apices continues as the functional apex of the stem (bifurcating again at each subsequent node), and the other functions as the apex of the peduncle. The peduncle first produces a bract and, then, a pedicel in the axil of the bract. Subsequent pedicels are each axillary to separate bracts. The pedicels, therefore, can be interpreted as ordinary lateral branches. However, because the bifurcations of the stem apex are not associated with subtending organs, the branching of the stem does not conform to expected monopodial or sympodial systems in the angiosperms. This suggests the possibility that each bifurcation of the stem apex is a true dichotomy. The anthoclad axis, thus, is a series of dichotomies. Although such a series may have been phylogenetically derived from a monopodial or sympodial ancestor, it is also possible that it may have been retained from a primitive, dichotomizing inflorescence.     

ULTRASTRUCTURE OF NON-ARTICULATED LATICIFERS IN MATURE EMBRYOS AND SEEDLINGS OF ASCLEPIAS SYRIACA L. (ASCLEPIADACEAE)

The protoplast of the non-articulated branched laticifer in the embryo and seedling of Asclepias syriaca L. was studied at the ultrastructural level and was found to differ from that of adjacent cell types. Embryonal laticifers possess numerous vesicles with electron-dense contents, but lack a large organized central vacuole. Plastids have few lamellae, possess phytoferritin, and accumulate small amounts of starch. Other organelles and membrane systems are similar to those in other cells. After germination, laticifers develop numerous elongated vacuoles by dilation of endoplasmic reticulum. Nuclei in laticifers within the hypocotyl of seedlings are highly lobed and possess dilated perinuclear spaces. Plastids and other organelles are similar to those observed in the protoplast of laticifers in the embryo. The latex or rubber component of the laticifer is not apparent in mature embryos of 72-hr seedlings.     

VARIATION IN THE POLLINARIA,ANTHERS, AND ALAR FISSURES OF COMMON MILKWEED (ASCLEPIAS SYRIACA L.)

We measured 10 characters of pollinaria, anthers, and alar fissures from the flowers of seven clones of common milkweed Asclepias syriaca L. to test whether significant differences existed among them. Strong among-clone differences existed for most of these characters. However, these differences were not correlated with each other, except for width of the alar fissure and width of the translator arm. Thus, these floral structures vary mostly in shape, rather than size per se. Few correlations among characters occurred at a within-clone level. Clones growing in the open and shade differed in half of the characters measured, but habitat did not substantially change the among-clone size patterns observed. The alar fissure–translator arm relationship might promote differential uptake of pollinia; however, A. syriaca does not exhibit a clear separation into donor and recipient clones.     

MATERNAL AND PATERNAL EFFECTS ON FOLLICLE PRODUCTION IN THE MILKWEED ASCLEPIAS SYRIACA (ASCLEPIADACEAE)

We measured follicle production from a diallel cross among ten clones of the common milkweed Asclepias syriaca, to assess the relative contributions of maternal and paternal parents. Specific parental combinations differed in the ability to set fruit, indicated by a significant nuclear specific effect accounting for 28% of the observed variance in follicle production. Several mechanisms might contribute to this effect, including shared incompatibility alleles and expression of zygotic genotypes. The nuclear general effect was not significant, however, suggesting a lack of additive genetic variation for offspring control of fruit maturation. Maternal effects also had an important effect on follicle production, as demonstrated by a significant reciprocal general effect (26% of the variance), almost entirely due to a large maternal component. The small reciprocal general variance component attributable to paternal effects, and nonsignificant reciprocal specific effect, indicating little maternal parent-zygote interaction, suggest that female choice through selective follicle maturation was not important in this experiment. The clones varied in proportion of reproductive output through female function, but a significant tradeoff between male and female success was not detected.     

ROLES OF POLLEN AND OVARY AGE IN FOLLICLE PRODUCTION OF THE COMMON MILKWEED ASCLEPIAS SYRIACA

Using cross-pollination experiments, pollinia viability of the common milkweed Asclepias syriaca L. (Asclepiadaceae) did not change significantly over the five-day life of the flowers, but stigma receptivity decreased over threefold over this period. During most of the season pollinators inserted over 90% of their pollinia into flowers during the first two days following anthesis. However, pollinators were scarce during the first few days of flowering, and they inserted over 60% of their pollinia into flowers three or more days old. This behavior potentially decreased outcrossing success and increased the relative male contribution of the flowers. Pollinator visitation patterns may play an important role in determining the functional sexuality of these plants.     

VARIATION AND EVOLUTION OF BREEDING SYSTEMS IN THE TURNERA ULMIFOLIA L. COMPLEX (TURNERACEAE)

The evolutionary and functional relationships among breeding systems and floral morphology were investigated in the Turnera ulmifolia complex. Predictions of a model of breeding system evolution among distylous and homostylous varieties were tested. Chromosome counts of 73 accessions revealed an association between breeding system and chromosome number. Diploid and tetraploid populations of five taxonomic varieties are distylous and self-incompatible, whereas hexaploid populations of three varieties are homostylous and self-compatible. The latter occur at different margins of the geographical range of the complex. Crossing studies and analyses of pollen and ovule fertility in F₁'s revealed that the three homostylous varieties are intersterile. To test the prediction that, homostylous varieties are long homostyles that have originated by crossing over within the distyly supergene, a crossing program was undertaken among distylous and homostylous plants. Residual incompatibility was observed in styles and pollen of each homostylous variety with patterns consistent with predictions of the cross-over model. The intersterility of hexaploid varieties suggests that long homostyly has arisen on at least three occasions in the complex by recombination within the supergene controlling distyly. Deviation from expected compatibility behavior occurs in populations of var. angustifolia that have the longest styles. These phenotypes displayed the greatest separation between anthers and stigmas (herkogamy) and set little seed in crosses with long- or short-styled plants. This suggests that they are derived from long homostyles with shorter length styles. It is proposed that selection for increased outcrossing has favored the evolution of herkogamy in long homostyles. Estimates of outcrossing rate in a distylous population using allozyme markers confirmed that dimorphic incompatibility enforces complete outcrossing. Significant genetic variation for floral traits likely to influence the mating system, such as stigma-anther separation, occurs within and among homostylous populations of var. angustifolia on Jamaica. Estimates of the mating system of families from a population with varying degrees of stigma-anther separation, using five isozyme loci, were heterogeneous and ranged from t = 0.04–0.79. Families exhibiting the largest mean stigma-anther separation have higher outcrossing rates than those with little separation.     

POLLINATION AND FRUIT-SET IN ASCLEPIAS: A REAPPRAISAL

Experimental pollinations of Asclepias tuberosa L., the first for this species and second large-scale effort for the genus, reveal trends toward local population differentiation. The species possesses a low level of self-compatibility, the first documented report for the genus. Analysis of flowers visited by natural pollinators in populations across the eastern half of the United States demonstrated a linear relationship between pollinia removal and pollinia insertion. Clarifications in older literature regarding the specificity of the pollination mechanism, effective levels of pollination in natural populations, and the performance of experimental manipulations are made, and it is concluded that fruit-set in Asclepias is regulated by the interaction of both mechanical aspects of pollination and physiological aspects of fertilization and fruit development. A model incorporating these restraints is seen to predict fairly well the observed fruit to flower ratio of about 1:100 in natural populations.     

镉在黑藻叶细胞中的亚显微定位分布及毒害效应分析

水生植物被认为能积累和浓缩重金属,本文以不同浓度Cd培养7天的黑藻为材料,研究了Cd的积累,亚细胞分布,亚显微定位,存在形态及对矿质营养吸收,光合速率和呼吸速率的影响,以揭示Cd的植物毒害机理.研究结果表明,随培养液中Cd浓度的增加,植株体内的Cd含量显著上升,富集系数为193-307;不同亚细胞组份中的Cd含量也都呈上升趋势,但分布显著不均,分配顺序依次为细胞壁＞可溶性部分＞细胞器,其中,积累在细胞壁中的含量所占比例呈下降趋势(61.66%-52.00%),而可溶性部分中则递增,细胞器部分结合的Cd含量所占比例相对恒定;硫化物-银法定位结果直观显示出Cd分布在细胞壁、叶绿体、细胞核和液泡中;逐步提取法分析表明受Cd毒害的黑藻体内Cd以不同化学形态存在,其中NaCl提取态占比例最大(55.05%),其他则较少,依次为FNaCl＞FHAc＞FWater＞FEthanol＞FHCl＞FResidue;Cd对黑藻的营养元素吸收也产生影响,主要是促进对Ca,Mn,Cu,Fe的吸收,降低对大量元素P,K的吸收;Cd对黑藻的光合速率和呼吸速率都有较强的抑制作用.结果表明Cd毒害与处理浓度间存在显著的剂量-效应关系;亚细胞分布与对细胞超微结构的损伤关系密切;Cd破坏了黑藻正常生理活动的结构基础和离子平衡,并造成功能紊乱.这些都是Cd对植物产生毒害的重要原因.     

EFFECTS OF VARIATION IN FLORAL MORPHOLOGY ON POLLINATION MECHANISMS IN ASCLEPIAS TUBEROSA L., BUTTERFLYWEED (ASCLEPIADACEAE)

Field observations revealed the presence of fruit-producing (fruiting) and non-fruit-producing (non-fruiting) plants in two south central Oklahoma populations of Asclepias tuberosa L. Comparative measurements of floral characteristics between fruiting and non-fruiting plants indicated that fruiting plants have significantly larger alar fissure widths and a greater percentage of intact pollinaria. The smaller alar fissure width on flowers of non-fruiting plants apparently reduces the probability of successful pollinia insertions in these plants. Greater numbers of observed pollinia insertions into fruiting-plant fissures appear to be the result of the increased likelihood that pollinia, which are significantly narrower than the fissures of fruiting plants, will be inserted into fruiting-plant fissures than those of non-fruiting plants. Apparently non-fruiting plants act primarily as pollinia donors while fruiting plants act primarily as pollinia recipients. These characteristics of butterflyweed tend to promote a dimorphic, allogamous type of pollination.     

FLORAL POLYMORPHY AND AMPHIMICTIC BREEDING SYSTEMS IN RUELLIA CAROLINIENSIS (ACANTHACEAE)

The taxonomically complex eastern U. S. species R. caroliniensis produces six functional or partially functional floral morphs. They are chasmogenes, reduced chasmogenes, morphologic protogynous chasmogenes, multistylous chasmogenes, cleistogenes, and semi-cleistogenes. The plants are uniformly self-compatible and generally produce normal fruits and seeds. The species has the morphological basis of a nearly balanced breeding system between allogamous and autogamous reproduction. Observations and comparisons on garden and greenhouse cultures suggest that R. caroliniensis is a seasonal cleistogamic species, and it is proposed that the different breeding systems are the chief causes of the difficult taxonomic problem in the species.