Nageiaceae—A New Gymnosperm Family

A new monotypic gymnosperm family, Nageiaceae D. Z. Fu, is separated from Podocarpaceae. It is characterized by having multinerved leaves without costae, and primitive shoot-like female reproductive organs (female strobili). The new family contains a single genus consisting of 2 sections, 5 species and is distributed along the western coast of the Pacific, from low coastal mountains of eastern and southern Asia to the Phillipines and Papua New Guinea. The first species in the Nageiaceae was described as an angiosperm, Myrica nagi Thunb. (1784), but it was soon recognized to be a gymnosperm belonging to a new genus, and was renamed as Nageia japonica Gaert. (1788). The generic name, Nageia, however, has seldom been used, and the members of Nageia have generally been treated as an isolated section of Podocarpus in the Podocarpaceae. When revising the Podocarpaceae, De Laubenfels (1969) established a new genus Decussocarpus based on Nageia, but several years later (1987) he revived the old generic name, Nageia. Page ( 1988,1990)considered Nageia to be a valid generic name and redefined it as a natural genus. The distinctive,broadly lanceolate, multinerved leaves (without costae) of Nageia are rather unusual in gymnosperms,only being similar to those of Agathis in the Araucariaceae, their leaves are also similar to each other in anatomy. For example, there are many single vascular bundles arranged parallelly, between which occur sclerenchyma cells in the mesophyll. Apparently,leaves in Nageia are rather similar both externally and internally to paleogymnosperm cordaitean leaves, and sclerenchyma cells found in Nageia might be the remains of straps between veins in cordaitean leaves. In addition to leaf characters, the large and nearly round pith of the young shoot in Nageia appears to be a reminiscent of the large pith in cordaitean stem. The female reproductive organs (female strobili ) in Nageia are shoot-like. The female strobilus has a sterile terminal bud, and several opposite or subopposite sterile scaly bracts on its axis; two opposite megasporophylls are found near the axis apex and both have an anatropous ovule which is almost entirely covered by the megasporophyll; a bract is partly adnate to the lower back of the megasporophyll;mature arillate seeds are 1-2 or occasionally 3 in number; the axis becomes woody when the seeds mature, but in some species (N. wallichiana) the upper part of the axis becomes fleshy (in the shape of a receptacle), in which no distinct boundary was found between the fleshy receptacle and the woody part, and both have the same scaly bracts or traces. Many characters in Nageia are distinctly different from those in Podocarpus. Leaves in the Podocarpaceae have distinct midribs; in Podocarpus, the reproductive organ, which was generally thought to be similar to that in Nageia, has no terminal bud, and its bract is entirely free from the lower back of the megasporophyll, the fleshy receptacle is derived from both the axis and the sterile bracts (except the lowest two), and the female strobilus at the seed stage has a secondary stalk. The multinerved leaf in Nageia can rarely be found in most of the living gymnosperms except in some rather isolated groups, such as Araucariaceae, Ephedraceae,Ginkgoaceae and Welwitschiaceae. Paleobotanical evidence shows that multinerved leaves have been found in all of the geological ages from the Paleozoic to the present, and such a shoot-like female reproductive organ as in Nageia was found in some paleogymnosperms. It is very difficult to determine the systematic positions of these fossil plants because of lacks adequate material of reproductive organs or even lack of complete vegetative organs. The vascular system and leaf characters of gymnosperms are considered to be very conservative, and the fact that the common leaf shape and venation exist in both fossil and living gymnosperms could imply that there exists a multinerved-leaved evolutionary line ( M-line ) in gymnosperms, which could be traced back to the paleogymnosperm cordaitean plants or even older ones with multinerved leaves. The different types of the female strobili (female reproductive organs) of living gymnosperms, regardless of having one or only several seeds without a typical cone or many seeds with a cone, might have been derived from shoot-like or spikelike female reproductive organs possessed by their common ancestor.The fossil eviden ce shows that the typical cone similar to those of living gymnosperms first appeared in the Jurassic, much later than the single-seeded fossil plant without cones. The seed fossil appeared in the late Devonian Period. It is very difficult to infer the relationships among living gymnosperms, which are hardly derived from one another. But an analysis of the strobili, including the axis structure and position, number, morphology and degree of adnation of the phyllomes on them, would be helpful to the study of their phylogeny. It is evident, therefore, that the gymnosperms with leaves having a midrib might also have a rather long evolutionary course,but no transition between the midrib and multinerved patterns of leaf venation has so far been found in both living and fossil plants. Finally, it is noteworthy that the Nageiaceae are distributed along the western coast of the Pacific, where many primitive representatives, both in gymnosperms and angiosperms, still survive. This would be advantageous to the consideration of Nageiaceae as a primitive representative, or a descendant of fhe paleogymnos-perm cordaitean plants.     

Seed Morphology, Anatomy and Ultrastructure ofPhyllocladusL. C. & A. Rich. ex Mirb. (Phyllocladaceae (Pilg.) Bessey) in Connection with the Generic System and Phylogeny

To analyse the status and phylogeny of the genusPhyllocladus,seedsof all seven species of the genus were studied. The complexof features of thePhyllocladusreproductive system point to theisolated position of the genus within the conifers. Relationswith Podocarpaceae s. l., Taxaceae s. l. and Cephalotaxaceaeappeared to be remote because a complex of features clearlydistinguishesPhyllocladusfrom the afore-mentioned taxa. We findit advisable to circumscribe the family Phyllocladaceae as Bessey(1907) did, and place it into the order Taxales Knobl. in Warming(1890). From investigations of the seeds it appears the genusPhyllocladusconsistsof seven species, forming five groups. There is a significanttendency for transformation of the female reproductive structureswithin the generic boundaries ofPhyllocladus—seeds, originallysolitary, tending to aggregate in various kinds of compact clusters.Copyright1999 Annals of Botany Company PhyllocladusL. C. & A. Rich, ex Mirb., seed anatomy, seed morphology, systematics, phylogenetic relationships, Phyllocladaceae, Podocarpaceae s. str., Acmopyleaceae, Nageiaceae, Austrotaxaceae, Amentotaxaceae, Torreyaceae, Taxaceae, s. str., Cephalotaxaceae, Taxales.     

The callistophytales (Pteridospermopsida): Reproductively sophisticated paleozoic gymnosperms

The Callistophytales nov. order is proposed for a small family of Pennsylvanian and possibly Permian pteridosperms. Plants conform to two species of Callistophyton and their reproductive organs. The small, eustelic sporophytes have a scrambling, shrub-like habit, profuse branching, adventitious roots and pinnately compound fern-like leaves with sphenopterid pinnules. Reproductive organs are borne on the abaxial surface of the pinnules, and consist of cardiocarpalean ovules and synangiate pollen organs with saccate pollen. Gametophyte development and reproductive biology conform closely to those of extant gymnosperms in the Coniferales. Anatomically preserved reproductive organs of the major groups of Paleozoic gymnosperms are compared, and a closer than currently recognized relationship of Paleozoic seed plants is proposed.     

Morphological "primary homology" and expression of AG-subfamily MADS-box genes in pines, podocarps, and yews

The morphological variation among reproductive organs of extant gymnosperms is remarkable, especially among conifers. Several hypotheses concerning morphological homology between various conifer reproductive organs have been put forward, in particular in relation to the pine ovuliferous scale. Here, we use the expression patterns of orthologs of the ABC-model MADS-box gene AGAMOUS (AG) for testing morphological homology hypotheses related to organs of the conifer female cone. To this end, we first developed a tailored 3'RACE procedure that allows reliable amplification of partial sequences highly similar to gymnosperm-derived members of the AG-subfamily of MADS-box genes. Expression patterns of two novel conifer AG orthologs cloned with this procedure-namely PodAG and TgAG, obtained from the podocarp Podocarpus reichei and the yew Taxus globosa, respectively-are then further characterized in the morphologically divergent female cones of these species. The expression patterns of PodAG and TgAG are compared with those of DAL2, a previously discovered Picea abies (Pinaceae) AG ortholog. By treating the expression patterns of DAL2, PodAG, and TgAG as character states mapped onto currently accepted cladogram topologies, we suggest that the epimatium-that is, the podocarp female cone organ previously postulated as a "modified" ovuliferous scale-and the canonical Pinaceae ovuliferous scale can be legitimally conceptualized as "primary homologs." Character state mapping for TgAG suggests in turn that the aril of Taxaceae should be considered as a different type of organ. This work demonstrates how the interaction between developmental-genetic data and formal cladistic theory could fruitfully contribute to gymnosperm systematics.     

Evolution of Reproductive Organs in Land Plants

LEAFY gene is the positive regulator of the MADS-box genes in flower primordia. The number of MADS-box genes presumably increased by gene duplications before the divergence of ferns and seed plants. Most MADS-box genes in ferns are expressed similarly in both vegetative and reproductive organs, while in gymnosperms, some MADS-box genes are specifically expressed in reproductive organs. This suggests that (1) the increase in the number of MADS-box genes and (2) the subsequent recruitment of some MADS-box genes as homeotic selector genes were important for the evolution of complex reproductive organs. The phylogenetic tree including both angiosperm and gymnosperm MADS-box genes indicates the loss of the A-function genes in the gymnosperm lineage, which is presumably related to the absence of perianths in extant gymnosperms. Comparison of expression patterns of orthologous MADS-box genes in angiosperms, Gnetales, and conifers supports the sister relationship of Gnetales and conifers over that of Gnetales and angiosperms predicted by phylogenetic trees based on amino acid and nucleotide sequences. Received 30 July 1999/ Accepted in revised form 9 September 1999     

Gymnosperm Orthologues of Class B Floral Homeotic Genes and Their Impact on Understanding Flower Origin

Class B floral homeotic genes play a key role in specifying the identity of male reproductive organs (stamens) and petals during the development of flowers. Recently, close relatives (orthologues) of these genes have been found in diverse gymnosperms, the sister group of the flowering plants (angiosperms). The fact that such genes have not been found so far, despite considerable efforts, in mosses, ferns or algae, has been taken as evidence to suggest that B genes originated 300–400 million years ago in a lineage that led to extant seed plants. Gymnosperms do not develop petals, and their male reproductive organs deviate considerably from angiosperm stamens. So what is the function of gymnosperm B genes? Recent experiments revealed that B genes from diverse extant gymnosperms are exclusively expressed in male reproductive organs (microsporophylls). At least for some of these genes it has been shown that they can partially substitute for the Arabidopsis B genes AP3 and PI in ectopic expression experiments, or even partially substitute these genes in different class B floral organ identity gene mutants. This functional complementation, however, is restricted to male organ development. These findings strongly suggest that gymnosperm and angiosperm B genes have highly related interaction partners and equivalent functions in the male organs of their different host species. It seems likely that in extant gymnosperms B genes have a function in specifying male reproductive organs. This function was probably established already in the most recent common ancestor of extant gymnosperms and angiosperms (seed plants) 300 million years ago and thus represents the ancestral function of seed plant B genes, from which other functions (e.g., in specifying petal identity) might have been derived. This suggests that the B gene function is part of an ancestral sex determination system in which B gene expression specifies male reproductive organ development, while the absence of B gene expression leads to the formation of female reproductive organs. Such a simple switch mechanism suggests that B genes might have played a central role during the origin of flowers. In the out-of-male and out-of-female hypotheses changes in B gene expression led to the origin of hermaphroditic flower precursors out of male or female gymnosperm reproductive cones, respectively. We compare these hypotheses with other recent molecular hypotheses on the origin of flowers, in which C/D and FLORICAULA/LEAFY-like genes is given a more prominent role, and we suggest how these hypotheses might be tested in the future.     

Characterization of the expression patterns of LEAFY/FLORICAULA and NEEDLY orthologs in female and male cones of the conifer genera Picea, Podocarpus, and Taxus: implications for current evo-devo hypotheses for gymnosperms

The identity of genes causally implicated in the development and evolutionary origin of reproductive characters in gymnosperms is largely unknown. Working within the framework of plant evolutionary developmental biology, here we have cloned, sequenced, performed phylogenetic analyses upon and tested the expression patterns of LEAFY/FLORICAULA and NEEDLY orthologs in reproductive structures from selected species of the conifer genera Picea, Podocarpus, and Taxus. Contrary to expectations based on previous assessments, expression of LFY/FLO and NLY in cones of these taxa was found to occur simultaneously in a single reproductive axis, initially overlapping but later in mutually exclusive primordia and/or groups of developing cells in both female and male structures. These observations directly affect the status of the "mostly male theory" for the origin of the angiosperm flower. On the other hand, comparative spatiotemporal patterns of the expression of these genes suggest a complex genetic regulatory network of cone development, as well as a scheme of functional divergence for LFY/FLO with respect to NLY homologs in gymnosperms, both with clear heterochronic aspects. Results presented in this study contribute to the understanding of the molecular-genetic basis of morphological evolution in conifer cones, and may aid in establishing a foundation for gymnosperm-specific, testable evo-devo hypotheses.     

On the Current Classification System of Paleozoic Seeds

This paper summarizes the history of classifications of Paleozoic seeds and revaluates the previous classification systems of Paleozoic detached seeds. The current status of studies on Paleozoic. gymnosperms: has been deteched seeds and whole fossil gymnosperms indicates that Seward’s classification system for Paleozoic seeds inadequate since all the seeds of Cardiocarpales in his system are not cordaitean female reproductive organs as Seward’s suggested. It is shown from investigations of whole fossil plants that the members of Cardiocarpales were derived from at least three different major groups of Paleozoic gymnosperms. Moreover, Meyen’s suggestion that the gymnosperms be classified based on symmetry of seeds has been little supported since all the fossil gymnosperms have not shown structurally preserved seeds and organic attachment. In order to relate detached seeds to whole fossil gymnosperms, the present author suggests that five families: Lagenostomaceae, Pachytestaceae, Callospermariaceae, Cryptospermaceae and Cardiocarpaceae be established for Paleozoic seeds and the Order Trigonocarpales be renamed as Pachytestales since the genus Trigonocarpus does not now include structurally preserved seeds. Thus, the five families may be considered either as subdivisions of the three orders of detached seeds: Lagenostomales, Pachytestales and Cardiocarpales, or as female reproductive organs of whole fossil gymnosperms of the five Permo-Carboniferous major groups: Lyginopteridales, Medullosales, Callistophytales Gigantopteridales and Cordaitales. A Key to Paleozoic seeds is provided as follows: A. Seeds with a cupule; integument thin, simple, deeply lobed and less differentiated;nucellus united to integument up to the base of pollen chamber; pollen chamber complex ................................. Lagenostomales, Lagenostomaceae A. Seeds without a cupule; integument thick, complex, unlobed and differentiated into several layers; nucellus free within integument except at the base; pollon chamber simple ................................................................................................ B B. Seeds radially symmetrical in shape; integumentary bundles present; nucellus bundles typical ................................................... Pachytestales, Pachytestaceae B. Seeds bilaterally symmetrical in shape; integumentary bundles present or absent; nucellus bundles often untypical .................................... C (Cardiocarpales) C. Bundles absent in integument; main bundle C-shaped in transverse section with a sclerenchyma bundle ............................................ Cryptospermaceae C. Bundles present in integument; main bundle not C-shaped in transverse section without a sclerenchyma bundle ......................................................... D D. Seeds very small with secretory cavities in integument; nucellus bundles limited in nucellus platform .......................................... Callospermariaceae D. Seeds large without secretory cavities in integument; nucellus bundles limited in nucellus platform or not ....................................... Cardiocarpaceae     

三叶木通花粉前纤维蛋白基因的克隆与表达

以三叶木通花蕾为材料,采用RT-PCR、3-′RACE方法克隆了三叶木通花粉前纤维蛋白基因,命名为Atf-Pro(GenBank登录号GQ478584)。结果表明:AtfPro的cDNA全长735 bp、阅读框393 bp、编码131个氨基酸,有1个342 bp的3′端非翻译区。预测分子量约为14.081 kD,等电点4.74。氨基酸和核苷酸序列的同源性分析发现,AtfPro基因属于植物花粉profilin基因家族的新成员。RT-PCR定性分析表明,AtfPro基因在三叶木通花蕾、花药、雌花花瓣和柱头组织中均有表达,但在幼叶、茎尖、根尖组织中低水平表达或不表达,生殖器官中的表达时期从花序分化发育开始到开花散粉结束。     

Sex determination in the monoecious species cucumber is confined to specific floral whorls

In unisexual flowers, sex is determined by the selective repression of growth or the abortion of either male or female reproductive organs. The mechanism by which this process is controlled in plants is still poorly understood. Because it is known that the identity of reproductive organs in plants is controlled by homeotic genes belonging to the MADS box gene family, we analyzed floral homeotic mutants from cucumber, a species that bears both male and female flowers on the same individual. To study the characteristics of sex determination in more detail, we produced mutants similar to class A and C homeotic mutants from well-characterized hermaphrodite species such as Arabidopsis by ectopically expressing and suppressing the cucumber gene CUCUMBER MADS1 (CUM1). The cucumber mutant green petals (gp) corresponds to the previously characterized B mutants from several species and appeared to be caused by a deletion of 15 amino acid residues in the coding region of the class B MADS box gene CUM26. These homeotic mutants reveal two important concepts that govern sex determination in cucumber. First, the arrest of either male or female organ development is dependent on their positions in the flower and is not associated with their sexual identity. Second, the data presented here strongly suggest that the class C homeotic function is required for the position-dependent arrest of reproductive organs.     

Comparisons among biomass allocation and spatial distribution patterns of some vine,pteridophyte, and gymnosperm shoots

The interspecific allometry of leaf, stem, and reproductive biomass distal to stem diameter was determined for a total of 12 angiosperm vine, gymnosperm, and pteridophyte species to compare allocation patterns to vegetative and reproductive shoot organs. The allometry of stem diameter in terms of the distance from shoot apices also was determined to quantify the manner in which vines, gymnosperms, and pteridophyte stems tapered along their length. The stems of vine species were found to weigh more than those of arborescent gymnosperm species distal to any point of equivalent stem diameter. Vine species also distribute more of their stem mass to shoot length as opposed to girth than gymnosperm species. Vine stems also supported proportionally larger leaf and reproductive biomass in comparison to gymnosperm stems of equivalent diameter, yet partitioned their total shoot biomass more or less equally between leaf and stem biomass in the same manner as the gymnosperm species examined. The allometry of vine as well as gymnosperm leaf biomass with respect to stem biomass appeared to be slightly anisometric and negative, suggesting that more massive stems had proportionally less leaf biomass than their smaller, less massive counterparts. Vine stems could be approximated as very slender cones; the shape and geometry of gymnosperm stems complied with those of stubby, truncated cones whose top diameter (for those examined), on the average, equaled 28% of the basal diameter. In general terms, the interspecific allometry of vines was most similar to that of pteridophytes. Collectively, these data refute the commonly held notion that vine stems are simply more slender than those of species with self-supporting stems.     

Anisakis physeteris: Amino acids in body tissues

The whole cuticle, perienteric fluid, and reproductive organs of Anisakis physeteris Baylis, 1923 from the sperm whale were analyzed for amino acid composition. Amino acid nitrogen per total nitrogen accounted for 96.2% in the cuticle. Corresponding values of TCA supernatant and precipitated fractions of perienteric fluid and reproductive organ were obtained. Proline, glycine, and arginine occurred abundantly in cuticle hydrolyzate. Lysine, glutamic acid, glycine, valine, leucine and histidine occounted for approximately 62% of total nitrogen in reproductive organ fractions. Differences were observed in the amounts of certain amino acids present in corresponding tissue of males and females as well as different tissue fractions from nematodes of the same sex, e.g., more serine in male cuticle than female, more proline in the protein of female reproductive organs than male; methionine present in cuticular protein and reproductive organ protein and nonprotein fractions but absent in perienteric fluid, and cystine in cuticular protein but not the other fractions. Untreated Anisakis hemolymph showed at least a 2-fold increase in ammonia in analyses made 2 and 10 days after bleeding. The relatively large ammonia content could have been in part the result of functional ornithine cycle and amino acid oxidase activity in vivo.     

Immuno-electron microscopic localization of estradiol receptor in cells of male and female reproductive and non-reproductive organs

Summary— The localization of estradiol receptor (ER) in various tissues and their distribution in sub-cellular compartments were studied by means of immunogold-electron microscopic methods using a site-directed polyclonal antibody developed against a peptide from the DNA binding site of ER. This method was used to determine the presence and localization of ER in tissues and cells of male and female reproductive and non-reproductive organs. In the female reproductive tract, endometrial cells and the cells of the corpus luteum were found to contain ER. In non-reproductive organs of both sexes the following cell types showed significant labeling: hepatocytes, epithelial duodenal cells, striated muscle fibers, cells of the proximal convoluted tubules of the kidney, lymphocytes, neurons, and adipose cells. Alveolar epithelial cells were studied only in female specimens and were labeled by the anti-ER. Prostatic and epididymal epithelial cells were found to be labeled in the male reproductive organs. In all these cells a higher density of label was found in the nucleus, especially in the space between the clumps of compact chromatin, as was previously found in epithelial endometrial cells. These results suggest that estradiol exerts its effects through a common nuclear mechanism in cells of male and female reproductive and non-reproductive organs.     

Ripe pollen structure and histochemistry of some gymnosperms

Some aspects of pollen cytology at dispersal were studied in 12 species of gymnosperms. The pollen grains differed in: 1. volume and cell number; 2. polarization of external structure and internal cell components; 3. wall thickness, especially of the intine, and the resulting percentage of cell volume with respect to total pollen grain volume; 4. stratification and chemical nature of the various intine layers; 5. nature and location of polysaccharide reserves; 6. morphological differences between the dry and hydrated states and phenomena related to hydration; 7. presence and site of orbicles. The various characters are compared and discussed in relation to the length of the reproductive cycle and the relations between the male gametophyte and its female counterpart.     

A Contribution to the Embryology of Ginkgo with a Discussion on the Affinity of the Ginkgoales

The ancestry of Ginkgoales or Ginkgophyta can be traced back to the Paleozoic. But today this order is represented only by a single species, Ginkgo biloba. Seward (1838) considered Ginkgo as one of the wonders of the world; it has persisted with little change until the present through a long succession of ages when the earth was inhabited by animals and plants for the most part far removed, in kind as in time, from their living descendents. Ginkgo is generally believed to be native to China, but so far it has not been found in wild state. A number of studies concerning the embryogeny of Ginkgo have been reported, on the basis of the materials accumulated during the past years and supplemented in 1978–1980, the embryogeny of Ginkgo is here described. Finally the phylogeny of Ginkgoales is discussed by comparing with the embryogeny of other groups of the living gymnosperms. Pollination usually takes place from the end of April to the first days of May and fertilization occurs around August 16–20 in the suburbs of Peking. Thus, the interval between pollination and fertilization is a few days less than four months. The embryo of Ginkgo is generally considered as suspensorless. In authors’ opinion, however, the somewhat elongated and much enlarged cells at the micropylar end of the embryo may be considered as the reduced suspensor cells though they are not the typical ones. There is no distinct demarcation between the proembryo and the young embryo in Ginkgo. In comparison with the embryogenesis in Coniferales, the tissue differentiation of the late embryo of Ginkgo is rather indistinct. Many authors such as Chamberlain (1935), Florin (1949), Delevoryas (1963), Sporne (1965) and others divide the gymnosperms into two major groups, Cycadophyta and Coniferophyta. From the point of view of morphological structure there are many significant common features shared by Ginkgoales and Coniferales. For example, Ginkgo possesses long shoot and short shoot while some conifers also have long and short shoots. The anatomical features of the stem of Ginkgo such as the well-developed secondary xylem, relatively small pith and the presence of bordered pits on the tracheids are also similar to those of the Coniferales. Not only Ginkgo has its characteristic leaf shape and the dichotomous venation but also they are quite different from the fronds of cycads. From the reproductive structure, on the other hand, Ginkgo and Cycadales are rather similar: both of them having one sulcate pollen grains, the pollen tube being of haustorial nature, the sperms being released from the base and not from the tip of the pollen tube, the presence of mulficiliate and rather large sperms, development of large female gametophyte bearing archegonia with exceptionally large eggs, more divisions of the free nuclear stage in the proembryo and less distinct differentiation of the ate embryo. All these features are primitive embryological characters. Thus, from the point of view of embryology the Ginkgo is close related to the Cycadales rather than to the Coniferales. Since Ginkgophyta are related to Cycadophyta in reproductive structures on one hand and similar to Coniferophyta in morphological and anatomical characters of the vegetative organs the Ginkgophyta are related to Cycadophyta in reproductive structures on one hand and similar to Coniferophyta in morphological and anatomical characters of the vegetative organs on the other hand, it indicates the interrelationship among these groups is clearly shown. Recently the discovery and studies on the progymnospermopsida (Beck, 1976) are worth notice, this fossil group contains the plants with certain characters of some conifers and certain characters of some cycads and this kind of plants bearing both homosporous and heterosporous forms are similar to ferns. They link the gymnosperms to the ferns. The present evidence, therefore, shows that the gymnosperms are very probably monophyletic and the progymnosperms might be the ancester of all the gymnosperms. The embryological characteristics supports the monophyletic origin of the gymnosperms.     

Indicators of sexual maturation and regression in the female cat flea, Ctenocephalides felis

Differences in the salivary glands, mesenteron epithelium and reproductive organs of female cat fleas, Ctenocephalides felis Bouché (Siphonaptera: Pulicidae), are related to the degree of reproductive maturation or regression. Contrary to previous ideas, blue bodies in the ovarioles are degenerate oocyte nuclei and their presence denotes failure of ripening oocytes to reach full maturity. A distinction between true corpora lutea and pseudo-corpora lutea is established, the presence of the former indicates successful oviposition, and of the latter, failure to complete maturation of eggs. Accurate indicators of sexual maturation and reproductive success are of potential value in assessing relative suitability of various hosts for a given flea species and therefore in assessing the degree of host specificity among fleas.