POLLEN MORPHOLOGY OF THE EUCHARIDEAE (AMARYLLIDACEAE)

Eucharis, Caliphruria, and Urceolina form a monophyletic group of petiolate-leaved, Neotropical Amaryllidaceae ecologically specialized to the understory of primary tropical rain forest below 2,000 m elevation. Pollen morphology of the three genera is surveyed. Pollen grains of all species of Eucharis, Caliphruria, and Urceolina are boat-shaped elliptic, monosulcate, heteropolar, and bilateral in symmetry. Exine sculpturing is semitectate-columellate and reticulate in all species examined. A transformation series in reticulum coarseness and pollen grain size is described. The large pollen grain with coarse reticulum of most Eucharis species is considered ancestral. The fine reticulation of Caliphruria is considered derived and the exine morphology of Urceolina is intermediate. Both of these genera have medium-sized pollen grains. Exine dimorphism common to all Urceolina, but rare in Eucharis and Caliphruria, may be symplesiomorphous among those taxa exhibiting this morphology. The three genera are largely uniform in pollen grain ultrastructure, with completely ektexinous exines. Pollen grain size in Eucharis is not closely correlated with style length. Several wide-ranging species show considerable intraspecific variation in pollen size. Parallelisms in pollen grain evolution among related tribes of Neotropical Amaryllidaceae are discussed.     

THE AURICULATE POLLEN GRAIN OF HYMENOCALLIS QUITOENSIS HERB. (AMARYLLIDACEAE) AND ITS SYSTEMATIC IMPLICATIONS

The pollen grain of Hymenocallis quitoensis Herb, is examined with scanning and transmission electron microscopy. The very large, monosulcate grain is characterized by a coarsely reticulate exine surface and solid, auriculate appendages at the equatorial facies. Surface morphology of the pollen grain of H. quitoensis is compared with other representative species of Hymenocallis and supports the inclusion of H. quitoensis within the genus. Meridional exine dimorphism in other species of Hymenocallis is interpreted as a reduction from the well-defined auriculae of H. quitoensis. Surface morphology of the pollen grain of Hymenocallis sects. Artema Traub and Elisena (Herb.) Traub exhibits divergence from that of sects. Hymenocallis and Ismene (Herb.) Traub. Phylogenetic and paleobotanical implications of the auriculate pollen grain are briefly discussed.     

POLLEN DEVELOPMENT IN OENOTHERA BIENNIS (ONAGRACEAE)

Development of the exine and viscin threads in Oenothera was studied by a combination of transmission electron microscopy and field emission scanning electron microscopy. Exine formation is initiated in the early tetrad stage by plate-like structures of preexine formed evenly around the microspore within a callosic wall. In the late tetrad stage, an endexine accumulates on white-lined lamellae underneath the preexine. After dissolution of the callosic wall, the preexine develops into beaded ektexine which is differentiated into an undulate tectum-like layer and columellae-like components. Interwoven fibrous strings connect the developing ektexine and the surface of the tapetal cells, and later develop into the viscin threads. These developmental processes imply that the columellae-like components are different in structure from the columellae of other angiosperms and that the formation of viscin threads is associated with the tapetal cells.     

THE INFLUENCE OF THE MATING SYSTEM ON SEED SIZE VARIATION IN CRINUM ERUBESCENS (AMARYLLIDACEAE)

We present evidence that extreme seed size variation in fruits of Crinum erubescens (range: 0.1 to 66.5 g per seed) occurs when mating pairs are inbred, either from selfing or biparental inbreeding. Several relatively uniform seeds of intermediate size are produced when pollen from several pollen donors is applied simultaneously to a flower. Selfed fruits and some fruits pollinated with a single pollen donor produce both large and small seeds, although selfed fruits produce fewer seeds than outcrossed fruit. These results are contrary to the hypothesis that variation in seed size is attributable to either pollen competition or differential allocation of maternal resource to seeds of different genotypes.     

POLLEN STUDIES IN THE GENUS CORDIA (BORAGINACEAE)

The pollen of 40 species of Cordia L. was studied by light microscopy and scanning and transmission electron microscopy. Three major types were found, 3-porate with a reticulate sexine, 3-colpate to 3-colporoidate with a striatoreticulate sexine, and 3-colpate to 3-colporate with a spinulose sexine. These pollen types are correlated with significant differences in the inflorescence, flower, and fruit structures found in the sporophyte. The following revisions are suggested and discussed: that Cordia sect. Varronia P. Br. be elevated to generic status; that Cordia sebestena L. and a few close allies be treated as generically distinct; and that Cordia myxa L. be conserved as the lectotype for the genus, thus continuing a wide application of the epithet Cordia.     

ANTHER AND POLLEN DEVELOPMENT IN RICE (ORYZA SATIVA)

Investigations of the growth of anthers and ontogeny of pollen grains of Oryza sativa (rice) IR-30 were undertaken for the purpose of 1) providing a set of growth measurements and 2) describing stable cytological features of anther and pollen development. Correlations exist between elongation of the floret and growth parameters of the anther such as its length, width, fresh and dry weights and cytological stage of pollen development. In the early ontogeny of the anther, hypodermal archesporial initials divide periclinally to form primary parietal cells and primary sporogenous cells. Each of the latter divides twice mitotically to generate four microspore mother cells, which undergo meiosis. The anther wall is formed by anticlinal and periclinal divisions of the primary parietal cells as well as of cells surrounding the primary sporogenous cells. Subsequent cytological features in the development of anther and pollen grains of rice have much in common with anther and pollen developmental biology of other members of Gramineae.     

DEVELOPMENT OF OMNIAPERTURATE POLLEN IN TRILLIUM KAMTSCHATICUM (LILIACEAE)

Ultrastructural changes during omniaperturate pollen development in Trillium kamtschaticum Pall, was examined using transmission electron microscopy. The pollen mother cells are not enveloped within a thick callosic wall. The microspores resulting from successive meiosis are divided by scanty deposition of callosic wall in the tetrad. A primexine/exine template is not recognizable within the tetrad during formation of exinous components. Preexinous globules, originating from vesicles in the callosic wall, accumulate electron-dense materials and develop into exinous globules. The preexinous globules have ca 10 nm wide contacts with tilted and invaginated plasma membrane of the microspore within the callosic wall. After dissolution of the callosic wall, the microspores separate and mitosis subsequently leads to the formation of a generative cell and vegetative cell encased in a loose aggregation of developing exinous globules. When the generative cell is at the pollen grain surface, the channeled zone is initiated at the opposite side of the microspore on the surface of the vegetative cell. Just before pollen maturity, a new layer develops under the channeled zone. Thus, development of the omniaperturate pollen grains of T. kamtschaticum involves some processes that are distinct from those of Canna and Heliconia and some that are similar.     

EFFECTS OF SOIL NITROGEN ON POLLEN PRODUCTION,POLLEN GRAIN SIZE,AND POLLEN PERFORMANCE IN CUCURBITA PEPO (CUCURBITACEAE)

To determine the effects of soil nitrogen on pollen production, pollen size, and pollen performance, two cultivars of zucchini (Cucurbita pepo) were grown under two nitrogen regimes in an experimental garden. The two cultivars were true breeding for alternative alleles for a one gene trait, ovary color. The soil nitrogen treatment had a significant effect on most measures of reproductive output through the female function. The nitrogen treatment did not affect the number of staminate flowers per plant but did have an effect on the number of pollen grains per staminate flower and the mean pollen grain size. A pollen mixture experiment revealed that pollen produced by plants in the high nitrogen treatment sired significantly more seeds than pollen from low nitrogen plants. Moreover, we found that the high nitrogen pollen sired even a greater percentage of seeds in the region of the fruit (ovary) previously shown to be fertilized by the fastest growing pollen tubes. Thus, the difference in the number of seeds sired by pollen from the two nitrogen treatments is due to differences in pollen performance. We conclude that spatial heterogeneity in soil nitrogen can influence the paternity of seeds in a plant population.     

POLLEN MORPHOLOGY AND EVOLUTION IN HEDYOTIS SUBGENUS EDRISIA (RUBIACEAE)

Primarily on the basis of aperture structure, 31 species of Hedyotis subg. Edrisia (Houstonia) are separable into 5 palynological groups. Group 1 is characterized by a simple os circumscribed by varying nexinous thickenings, groups 2–4 combine this os with a crassimarginate one to form a compound os, while group 5 is known only with the crassimarginate os. The first type is considered primitive, the compound os specialized and more advanced, and the third type reduced and highly advanced. Other characteristics of the pollen, including size and shape, aperture number, thickness of sexine and nexine, and reticulum, are discussed in relation to the apertures, and their probable primitive and advanced expressions are outlined. The data agree with the phyletic trends found in the sporophyte on the basis of results from morphology, chromosome number and size, and distribution, and they support and add to an earlier phylogenetic scheme proposed for the subgenus. The evidence from palynology also supports the treatment of Hedyotis and Houstonia as congeneric.     

POLLEN MORPHOLOGY IN THE GENERA MONTTEA AND MELOSPERMA (SCROPHULARIACEAE)

Pollen of the scrophulariaceous genera Melosperma and Monttea was studied with light and scanning electron microscopy. Both have single, radiosymmetric, isopolar, trizoniaperturate pollen grains with usually ruptured colpus membranes, tectate-perforate to semitectate exines with a columellate interstitium and pitted or perforate nexines. Intergeneric differences in pollen morphology are established based on the differential correlation of sculpturing characters. Comparisons of the pollen of Melosperma and Monttea with that of the tribe Minuleae sensu stricto reveals no difference in the case of Melosperma and the existence of intermediate morphologies and “reticulate” similarities in the case of Monttea; the taxonomic significance of this morphological intergradation is considered. The transfer of Oxycladus aphyllus to Monttea is consistent with the pollen evidence, but proposed affinities between Monttea and tribe Antonieae of the Loganiaceae are not supported.     

EARLY CRETACEOUS TETRADS,ZONASULCULATE POLLEN,AND WINTERACEAE. I. TAXONOMY,MORPHOLOGY, AND ULTRASTRUCTURE

Ulcerate pollen tetrads from the late Barremian-early Aptian of Gabon, named Walkeripollis gabonensis gen. et sp. nov., resemble pollen of extant Winteraceae but have finer sculpture and a weakly calymmate tectum, like tetrads reported from the late Aptian-Albian of Israel. Scanning and transmission electron microscopy reveal additional winteraceous features (tall muri, short columellae, ring of endexine around the ulcus), plus segmented muri recalling the reticulate, zonasulculate to inaperturate genus Afropollis, which is abundant in the Aptian-Albian of Northern Gondwana. Afropollis also resembles Winteraceae in having tall muri and short columellae, but it has a thicker endexine. The new zonasulculate genus Schrankipollis, including S. mawhoubensis (Schrank) comb. nov. from the Aptian of Egypt and S. microreticulatus (Brenner) comb. nov. from the Potomac Group of Maryland, resembles Afropollis in structure of its muri but differs in its elliptical shape, finer reticulum, and restricted endexine.     

POLLEN TUBE GROWTH AND INBREEDING DEPRESSION IN AMSINCKIA GRANDIFLORA (BORAGINACEAE)

To discriminate between the roles of differential pollen tube growth and inbreeding depression in causing reduced seed production following self-pollination of distylous Amsinckia grandiflora (Boraginaceae), callose plug and pollen tube production were compared following cross- and self-pollinations of A. grandiflora. Pin (long-styled morph) × thrum (short-styled morph) crosses yielded more pollen tubes than pin × pin crosses or pin self-pollinations; no difference could be detected for the pin × pin or pin self-pollination categories. Using thrums as the female parent, no significant differences were detected in any pollination category. These results indicate that reduced seed production following self-fertilization of A. grandiflora is likely to be the result of inbreeding depression rather than inhibition of self pollen tubes.     

CHROMOSOME CYTOLOGY OF EUCHARIS,CALIPHRURIA, AND URCEOLINA (AMARYLLIDACEAE)

Karyotypes are presented for 19 taxa of Eucharis, Caliphruria, and Urceolina, a monophyletic group of Andean Amaryllidaceae within “infrafamily” Pancratioidinae Traub. All three genera are characterized by 2n = 46, the most common somatic number occurring in the “infrafamily.” Incidences of polyploidy are low. Only two tetraploid (2n = 92) species of Eucharis are so far known, E. bouchei and E. bonplandii, the northernmost species of E. subg. Eucharis. The 2n = 68 karyotype of E. amazonica is interpreted as triploid-derived (3x – 1). Chromosomal heteromorphism is reported for C. subedentata. Karyotype data is analyzed with principal component analysis and unweighted pairgroup cluster analysis. In a number of cases, phenetic relationships among the karyotypes correlate with phenetic and cladistic relationships based on morphological data. Karyotype evolution among the three genera is discussed in the context of classical theories of karyotypic symmetry. Stability of chromosome number in Eucharis and related genera suggests that chromosomal evolution has proceeded via nonreciprocal interchanges between chromosomes and infrachromosomal structural change. In at least one case (E. bakeriana), rapid sympatric speciation may have been vectored by chromosomal change.