Polarity and competition between megaspores in the ovule ofOenothera hybrids

New data on the development of polarity in the ovules during megasporogenesis and early stages of embryo sac development inOenothera-hybrids are presented. It is confirmed that allOe. hookeri-hybrids show a strong tendency to form heteropolar tetrads, with the micropylar megaspore developing into an embryo sac. This preference is seen in the delay of the second meiotic division on the chalazal side, the absence of callose in the lateral wall of the micropylar megaspore, and the accumulation of starch in this megaspore. However, homopolar tetrads, chalazal preference, and ovules with two developing embryo sacs are also observed with considerable frequency. Quantitative data on the frequency of the different developmental types are compared with earlier genetic results about competition in the haplophase. There is sufficiently good agreement to support the hypothesis ofRenner that there is a correlation between the developmental processes in the megaspore tetrad and the genetic phenomena of competition in the haplophase.     

水稻亚种间杂种小穗败育的细胞学基础

本文对普通栽培稻不同品种类型间杂种小穗败育的细胞学基础及雌性败育的过程进行了研究,结果表明:(1)引起杂种小穗败育的原因有胚囊败育、花粉败育、开花时花药不开裂和雌雄异熟。其中胚囊败育而丧失受精能力是引起低结实率的最重要的因素,开花时花药不开裂和雌雄异熟在一定程度上形成了雌雄性细胞时间和空间的隔离屏障。(2)杂种植株的所有大孢子母细胞都能进行正常的减数分裂,形成四个大孢子,败育主要发生在靠近合点端的功能大孢子分化形成胚囊的早期,有的功能大孢子在进行第一次有丝分裂前便萎缩解体,多数走向败育的功能大孢子能完成一次或二次有丝分裂,形成二核或四核败育胚囊。败育的共同特征是无液泡的分化,细胞质少或退化,在败育胚囊残迹部位,解体的珠心细胞和萎缩的胚囊残渍混杂垛叠。已受精的杂种子房没有观察到胚及胚乳发育的异常。籼粳杂种胚囊败育频率较高。     

Different patterns of callose wall formation during megasporogenesis in two species ofOenothera (Onagraceae)

The homozygousOenothera hookeri Torr. etGray shows the typical pattern ofOnagraceae with ± callose on the external walls of megaspore mother cells and tetrads. Megasporogenesis is heteropolar, and the micropylar megaspore is the mother-cell of the 4-celled embryo sac. The complex-heterozygousOenothera biennis L. during megasporogenesis generally has callose not on the external cell walls but only on the transversal walls of the tetrad. In 95% of the ovules both the external chalazal and the micropylar megaspores develop to embryo sac mother-cells. Megasporogenesis is homopolar, and competition between two developing embryo sacs for nutrition in the ovule occurs. The embryo sac with the stronger genotype wins the race against the other one. Polarity phenomena during ontogeny of the female gametophyte are related to nutritional supply and hormonal induction from the ovule. The introduction of a developmental-physiological point of view into the discussion about the evolution of the embryo sac inOnagraceae is therefore justified.Stipendiatin der Alexander von Humboldt-Stiftung 1974/76.     

Somatic embryogenesis in soybean via somatic embryo cycling

Summary The objectives of the present research were: a) to develop an efficient soybean embryogenic regeneration system characterized by a high frequency of explant response and a large number of somatic embryos per explant; b) to evaluate the factors affecting somatic embryogenesis via somatic embryo cycling; and c) to identify the origin of somatic embryos in the system. A highly improved and efficient system for soybean somatic embryogenesis was established using somatic embryo cotyledons and somatic embryo hypocotyl/radicle explants plated on α-naphthaleneacetic acid (NAA) or 2,4-dichlorophenoxyacetic acid (2,4-D) supplemented MS basal media. The system included somatic embryo cycling between liquid and solid medium and it consistently gave rise to a much higher frequency of explant response and a larger number of embryos per responding explant than those obtained from zygotic cotyledon explant tissues. Genotype, differences were observed for response in some of the treatments with cv “Fayette” being more responsive than “J103”. Histological studies revealed that somatic embryos induced in the somatic embryo cycling system originated almost exclusively from epidermal cells on both 2,4-D and NAA inductive media. The cells of the epidermis proliferated to produce somatic embryos directly without an intervening callus phase. A single-cell origin of somatic embryos was observed in cultures on a 40 mg/liter 2,4-D treatment. A large number of responding cells in the epidermis was also observed in the 10 mg/liter NAA treatment. The single-cell origin of somatic embryos from epidermal layers of the explant tissues should facilitate development of an efficient transformation system for soybean.     

水稻PDER品系的特点和二倍体孢子生殖胚囊的形成和发育

对水稻（ＯｒｙｚａｓａｔｉｖａＬ．）早发生胚ＰＤＥＲ（ｐｒｅ－ｄｅｖｅｌｏｐｅｄｅｍｂｒｙｏｏｆｒｉｃｅ）品系的特点和细胞胚胎学研究表明,ＰＤＥＲ是二倍体植物２ｎ＝２４,约有５０％胚囊的卵细胞未经受精能自行发育形成胚,成熟种子的萌发和生长速度较常规正常水稻快。ＰＤＥＲ的大孢子母细胞经有丝分裂产生未减数的胚囊,即无融合生殖中的二倍体孢子生殖类型。在胚囊形成和发育过程中有如下几个特点：（１）孢原细胞至大孢子母细胞分裂前的过渡期持续时间较长,孢原细胞和大孢子母细胞的细胞质比周围的珠心细胞质稀淡。（２）大孢子母细胞经二次有丝分裂后形成直线排列的三个细胞（三分体）,珠孔端的两个解体,合点端的一个发育为功能细胞,有少数胚囊的三个细胞全部解体形成败育胚囊。（３）功能细胞经三次连续核分裂形成具八核七个细胞的成熟胚囊,它的结构与常规正常水稻基本相同,但助细胞呈长形而没有回抱着卵细胞。     

Spores of microorganisms

Calcium-dipicolinate (Ca-DPA)-rich and Ca-DPA-deficientBacillus cereus spores were incubated in a synthetic medium with germination stimulants and in bactopeptone medium with a fairly high calcium ion concentration. In the complex medium the germination of Ca-DPA-rich spores was completely blocked at a concentration of 0.5m CaCl₂, whereas the complete blockage of germination in the synthetic medium required higher concentrations (0.6–0.8m) of calcium chloride. Ca-DPA-deficient spores germinated more slowly and less completely in the synthetic medium than in the bactopeptone medium. The germination of these spores took place, however, even at higher calcium ion concentrations (0.6–0.8m). On the contrary, lower calcium chloride concentrations (0.1–0.4m) accelerated the germination of these spores in the synthetic medium and the final percentage of phase-dark and stainable spores was higher. “H-forms” of the Ca-DPA-rich and Ca-DPA-deficient spores prepared by acid titration germinated in both media. The germination of the latter spores being slower and proceeding less completely. “H-forms” germinated completely or partially in media with a high concentration of calcium chloride. The percentage of germinated spores, however, was strongly influenced by the concentration of this cation, especially the “H-forms” of Ca-DPA-deficient spores. Moreover, the germination of Ca-DPA-deficient spores in this medium was affected by the length of previous storage and, in the case of “H-forms” by the pH at which they were titrated. It was assumed that the increased permeability of calcium into the calciumundersaturated spore periphery in Ca-DPA-deficient and in “H-forms” of spores of both types co-determines (in the presence of germinants) the germinability of bacterial spores.     

Study of aberrant morphologies and lack of conversion of somatic embryos of chickpea (Cicer arietinum L.)

Summary Somatic embryos which originated from mature embryo axes of the chickpea (Cicer arietinum L.) showed varied morphologies. Embryos were classified based on shape of the embryo and number of cotyledons. “Normal” (zygotic-like) embryos were bipolar structures with two cotyledons and a well-developed shoot and root apical meristem, whereas “aberrant” embryos were horn-shaped, had single and multiple cotyledons, and were fasciated. Histological examination revealed the absence of a shoot apical meristem in horn-shaped embryos. Fasciated embryos showed diaxial fusion of two embryos. Secondary embryogenesis was also observed, in which the embryos emerged from the hypocotyl and cotyledonary region of the primary somatic embryo. This report documents the absence of an apical meristem as a vital factor in the lack of conversion of aberrant somatic embryos.     

Megasporogenesis,Microsporogenesis and Development of Gametophytes in Eleutherococcus senticosus (Araliaceae)

This paper describes megasporogenesis, microsporogenesis, and development of female and male gametophytes in Eleutherococcus senticosus. The main results are as follows: Flowers of E. senticosus are epigynous, pentamerous. Anthers are 4 -microsporangiate. An ovary has 5 loculi. Each ovary loculus has 2 ovules: the upper ovule and the lower ovule. The upper one is orthotropous and degenerates after the formation of archesporial cell, while the lower one is anatropous, unitegmic and crassinucellar, and able to continue developing. In male plants, microsporogenesis and development of male gametophytes took place in regular way, but a series of abnormal phenomena were found in megasporogenesis and development of female gametophytes. The microspore mother cells gave rise to tetrahedral tetrads by meiosis. Cytokinesis was of the simultaneous type. The mature pollen was 3-celled and shed singly. The anther wall formation belonged to the dicotyledonous type. At the stage of microspore mother cell, the anther wall consisted of four layers, i.e. epidermis, endothecium, middle layer, and tapetum. The tapetum was of glandular type and its most cells were binucleate. When microspores were at the uninucleate stage, the tapetum began to degenerate in situ. When microspores developed into 3-celled pollen grains, the tapetum had fully degenerates. In the lower ovule of male flower, the megaspore mother cell gave rise to a linear or “T” -shaped tetrad. In some cases, a new archesporial cell over the tetrad or two tetrads parallel or in a series were observed. Furthermore, the position of functional megaspore was variable; any one or two megaspores might be functional, or one megaspore gave rise to a uninucleate embryo sac, but two other megaspores also had a potentiality of developing into the embryo sac. In generally, on the day when flowers opened, female gametophytes contained only 4 cells: a central cell, two irregular synergids and one unusual egg cell. In female plants, microspore mother cells and secondary sporogenous cells were observed. But at the stage of secondary sporogenous cell, the newly differentiated tapetum took the appearance of degeneration. Later, during the whole stage of meiosis, the trace of degenerative tapetum could be seen. At last, the microsporangium degenerated and no tetrad formed. On the blossom day, all anthers shriveled without pollen grains. In female flowers, megasporogenesis and development of female gametophytes were normal: the tetrad of megaspores was linear or “T”-shaped; the chalazal megaspore was usually functional; the development of embryo sac was of the Polygonum type. On the blossom day, most embryo sacs consisted of 7 cells with 8 nuclei or 7 cells with 7 nuclei; but the egg apparatus was not fully developed. In hermaphroditic plants, microsporogenesis was normal but the development of male gametophytes was partially abnormal. When the hermaphroditic flowers blossomed, there were more or less empty pollen grains in the microsporangium and these pollen grains were quite different in size. The development of most gynoecia was normal but numerous abnormal embryo sacs could be seen. On the blossom day, female gametophytes were mainly 7-celled with 8-nuclei or with 7-nuclei or 4-celled with antipodal cells degenerated; the egg apparatus wasnot fully developed either.     

Ultrastructural characterization of apospory in Panicum maximum

The nucellar ultrastructure of apomictic Panicum maximum was analyzed during the meiocytic stage and during aposporous embryo sac formation. At pachytene the megameiocyte shows a random cell organelle distribution and sometimes only an incomplete micropylar callose wall. The chalazal nucellar cells are meristematic until the tetrad stage. They can turn into initial cells of aposporous embryo sacs. The aposporous initials can be recognized by their increased cell size, large nucleus, and the presence of many vesicles. The cell wall is thin with few plasmodesmata. If only a sexual embryo sac is formed, the nucellar cells retain their meristematic character. The aposporous initial cell is somewhat comparable to a vacuolated functional megaspore. It shows large vacuoles around the central nucleus and is surrounded by a thick cell wall without plasmodesmata. In the mature aposporous embryo sac the structure of the cells of the egg apparatus is similar to each other. In the chalazal part of the egg apparatus the cell walls are thin and do not hamper the transfer of sperm cells. Structural and functional aspects of nucellar cell differentiation and aposporous and sexual embryo sac development are discussed.     

Detection of Concealed “Illegitimate” Nuclei in Tetrad Analysis of the Diploid Progeny of Heterokaryons in Saccharomyces cerevisiae

In this work, the studies on the previously detected phenomenon of concealed heterokaryosis in Saccharomyces cerevisiaewere continued. In genetic and Southern blotting experiments, one of the nuclei in the heterokaryon was shown to be active (capable of division and ensuring the corresponding cell phenotype), whereas the other was not expressed until the heterokaryotic clone was transferred to the medium selective for this concealed nucleus. Moreover, the concealed nucleus was able to assume the active state after fusion with the second parental nucleus. It was analyzed whether the nuclei with new marker combinations occurring in meiosis can behave as exceptional nuclei. Tetrad analysis of hybrids carrying the kar1mutation in their nuclei revealed the relatively high percentage of exceptional tetrads (more than 10%). One spore in these tetrads usually formed diploid cells capable of sporulation. The presented data of genetic and molecular biological studies testify in favor of the assumption that abnormal spores contain two nuclei, which form an illegitimate hybrid after fusion. An extraneous nucleus (termed x) has usually a genotype close to that of one of the spores in this tetrad. Thus, it was assumed that the additional DNA replication round occurs in the absence of cell division during one of meiotic divisions. Results of cytological analysis conducted by the method of specific DNA staining confirmed the existence of exceptional tetrads, one spore of which contains two nuclei.     

高山红景天胚胎学研究

高山红景天(Rhodiola sachalinensis A.Bor.)具8个雄蕊,每个雄蕊有4个花粉囊。小孢子母细胞减数分裂时,胞质分裂为同时型。形成的四分体为四面体形。花药壁由表皮、药室内壁、二层中层和绒毡层五层细胞组成,其发育方式为基本型。腺质型绒毡层,有些绒毡层细胞分裂形成不规则双层,少数细胞双核。二细胞型花粉。雌蕊由4心皮组成。边缘胎座,倒生胚珠,双珠被,厚珠心,胚珠发育中形成珠心喙。大孢子四分体线形或T -形,合点大孢子具功能。胚囊发育为蓼型。成熟胚囊中,卵细胞核、助细胞核均位于细胞的合点端,珠孔端具液泡;极核融合为次生核,并位于卵细胞合点端附近; 3个反足细胞退化。双受精属于有丝分裂前配子融合类型。胚的发育为石竹型;基细胞侵入珠孔端,形成囊状吸器。细胞型胚乳;初生胚乳核分裂形成两个细胞,其珠孔端的细胞发育成胚乳本体,合点端的细胞直接发育成具一单核的合点吸器。     

An <Emphasis Type="Italic">Hieracium</Emphasis> mutant, <Emphasis Type="Italic">loss of</Emphasis><Emphasis Type="Italic">apomeiosis 1</Emphasis> (<Emphasis Type="Italic">loa1</Emphasis>) is defective in the initiation of apomixis

Asexual seed formation (apomixis) in Hieracium aurantiacum occurs by mitotic embryo sac formation without prior meiosis in ovules (apomeiosis), followed by fertilization-independent embryo and endosperm development. Sexual reproduction begins first in Hieracium ovules with megaspore mother cell (MMC) formation. Apomixis initiates with the enlargement of somatic cells, termed aposporous initial (AI) cells, near sexual cells. AI cells grow towards sexually programmed cells undergoing meiosis, which degrade as the dividing nuclei of AIs obscure and displace them. Following Agrobacterium-mediated transformation of an aneuploid Hieracium aurantiacum apomict, a somaclonal mutant designated “loss of apomeiosis 1” (loa1) was recovered, which had significantly lost the ability to form apomictic seed. Maternal apomictic progeny were rare and low levels of germinable seedlings were primarily derived from meiotically derived eggs. Cytological analysis revealed defects in AI formation and function in loa1. Somatic cells enlarged some distance away from sexual cells and unlike AI cells, these expanded away from sexual cells without nuclear division. Surprisingly, many accumulated callose in the walls, a marker associated with meiotically specified cells. These defective AI (DAI) cells only had partial sexual identity as they failed to express a marker reflecting entry to meiosis that was easily detected in MMCs and they ultimately degraded. DAI cell formation did not lead to a compensatory increase in functional sexual embryo sacs, as collapse of meiotic embryo sacs was prevalent in the aneuploid somaclonal mutant. Positional cues that are important for AI cell differentiation, growth and fate may have been disrupted in the loa1 mutant and this is discussed. The authors Takashi Okada, Andrew S. Catanach and Susan D. Johnson made equal contributions to the data.     

Comparative embryology and mammalian cloning

A hypothesis has been advanced that logically combines “contradictory” facts concerning the early mammalian development and shows a natural relationship between the embryos developing from a fertilized ovum and from cells of the inner cell mass of blastocyst. When studying the theoretical questions of cloning, it is necessary to take into consideration the peculiarities of prenatal mammalian ontogeny, which make themselves evident upon comparison with other animals. The absence of yolk in the mammalian ovum defines sharp differences in the early development between mammals and other Amniota. The complete asynchronous cleavage results in the formation of morula followed by blastocyst, which hatches from zona pellucida and is implanted into the uterus tissue. This fact allows us to consider the blastocyst as a mammalian larva, which is fed owing to the maternal organism. It is known that, in the body of a larva (blastocyst), a new embryo develops from some somatic cells. This process is known as polyembryony, which is typical of the development of some parasitic insects. Polyembryony in turn is a variant of somatic embryogenesis, which is a form of asexual reproduction. Thus, the two different embryos, “conceptus” and “embryo proper,” have different origins: the first forms by the sexual way and the second, by the asexual way. Investigation of the mechanisms of somatic embryogenesis in mammals will help us to find conditions necessary for full reprogramming of donor somatic nuclei and provide for successful development of reconstructed embryos.     

Development of Ovules and Embryo sacs in Ostrya virginiana (Betulaceae) and Its Systematic Significance

The development of ovules and embryo sacs in Ostrya virginiana was studied for the first time. Most ovaries had two ovules which were anatropous, unitegmic and crassinucellate. The ovule usually possessed several archesporial ceils which divided periclinally into the upper parietal cell and the lower sporogenous cell. The sporogenous cell functioned directly as megaspore mother cell. The tetrad of megaspores was linear in arrangement, and every megaspore might be functional. One ovule often contained 2- 6 embryo sacs and the embryo sac belongs to Polygonum type. It can be concluded from the present data that all ovules among the genera of the Betulaceae are unitegrnic. There are more groups with the phenomenon of multiple embryo sacs in anemophic plants such as Betulaceae, Casuarinaceae, Graminae, Jnglandaceae, Myricaceae, Simaroubaceae, Ulmaceae, than in entomophilous plants. Multiple embryo sacs also occur among some parasitic plants and saprophytes, e.g. Orobanchaceae, Cassytha in Lauraceae, Cuscuta in Convolvulaceae and Utricularia in Lentibulariaceae. It may be inferred that the characteristic of multiple embryo sacs be an evolutionary adaptation of those plants with lower pollination rate to increase the rate of fertilization. Finally, a comparison of embryological characters among the genera of the Betulaceae shows that the family is of a number of common embryological characters, such as multicellular archesporium, multiple embryo sacs in one ovule, and a long interval between pollination and fertilization. The diversity and systematic significance of several embryological characters among the “higher” hamamelid families are also discussed. It is still hard to explain the phy-logenetic relationships among those families clearly only with.     

Embryological Studies in Glycyrrhiza uralensis Fisch.

This paper reports the studies of overall embryology of Glycyrrhiza uralensis Fisch. Development of the anther wall follows the dicotyledonous type. The cytokinesis of the microspore mother cell in meiosis is of simultaneous type. The arrangement of microspores in tetrad is tetrahedral, isobilateral and decussate. Microspores have various types of abortive to development. Mature pollen grain is of the 2-celled type. The ovule is bitegminous, crassinucellate and campylotropous. The megaspore mother cell gives rise to unequal dyad and then linear tetrad. The chalazal megaspore, the second or the third megaspore towards the micropylar end are functional megaspore. The development of the embryo sac conforms to the Polygonum type. Mature embryo sac has various types of variation. The fertilization belongs to the premitotic type of syngamy. The development of most embryoes belongs to the Onagrad type. The development of the endosperm belongs to the nuclear type and the endosperm near the chalazal end develops into haustorium.     

High-frequency plant regeneration through cyclic secondary somatic embryogenesis in black pepper (Piper nigrum L.)

A high-frequency plantlet regeneration protocol was developed for black pepper (Piper nigrum L.) through cyclic secondary somatic embryogenesis. Secondary embryos formed from the radicular end of the primary somatic embryos which were originally derived from micropylar tissues of germinating seeds on growth regulator-free SH medium in the absence of light. The process of secondary embryogenesis continued in a cyclic manner from the root pole of newly formed embryos resulting in clumps of somatic embryos. Strength of the medium and sucrose concentration influenced the process of secondary embryogenesis and fresh weight of somatic embryo clumps. Full-strength SH medium supplemented with 1.5% sucrose produced significantly higher fresh weight and numbers of secondary somatic embryos while 3.0 and 4.5% sucrose in the medium favored further development of proliferated embryos into plantlets. Ontogeny of secondary embryos was established by histological analysis. Secondary embryogenic potential was influenced by the developmental stage of the explanted somatic embryo and stages up to “torpedo” were more suitable. A single-flask system was standardized for proliferation, maturation, germination and conversion of secondary somatic embryos in suspension cultures. The system of cyclic secondary somatic embryogenesis in black pepper described here represents a permanent source of embryogenic material that can be used for genetic manipulations of this crop species.     

Basidiocarp development inCoprinus macrorhizus

The dikaryon #5026 (A2B2)+#5132 (A7B7) of the basidiomyceteCoprinus macrorhizus was treated with ultraviolet light (UV),N-methyl-N′-nitro-N-nitrosoguanidine (NG), and 5-bromouracil (BU) in order to induced developmental variations in fruiting. Among a total of 11,383 isolates, 742 were homokaryons. Fruiting test was made with the remaining 10,641 dikaryotic isolates, of which 1,594 showed abnormality in basidiocarp development. Both UV and NG were very efficient in inducing such variations, but BU was not. The induced variations were classified into seven basic types as follows: 1) “knotless”, in which hyphal knots, the first sign of fruiting, do not differentiate; 2) “primordiumless,” in which hyphal knots are formed, but no further development occurs; 3) “maturationles”, in which primordia are formed, but the maturation of primordia into adult fruit bodies does not occur; 4) “elongationless”, in which the elongation of stipe is blocked, and a basidiocarp with a very short stipe is produced; 5) “expansionless,” in which pilei do not expand normally; 6) “sporeless,” in which the formation of basidiospores is blocked and albinic pilei bearing none or only a small amount of spores are produced; and 7) “autolysisless,” in which the autolysis of pilei does not appear to occur. It has been noticed that the four steps of maturation, i.e. stipe elongation, pileus expansion, spore formation, and perhaps pileus autolysis, can proceed independently. Compound types of variations such as “elongation-expansionless,” “elongation-sporeless,” “expansion-sporeless,” and “elongation-expansion-sporeless” were also induced. UV treatment induced maturationless at the highest rate, while NG treatment sporeless. Contributions from the Division of Plant Morphology, Department of Biology, Faculty of Science, Okayama University. No. 117.     

Evolutionary ontogenetic aspects of pathogenetics of chronic human diseases

This article is a review of scientific publications, in which issues of pathogenetics of multifactorial diseases (MFDs) are considered from the viewpoint of evolution and ontogeny. Concepts explaining significance of evolutionary processes in the formation of genetic architecture of human chronic diseases (“thrifty” genomes and phenotypes, “drifty genes,” decanalization) are analyzed. The roles of natural selection and genetic drift in the formation of hereditary diversity of genes for susceptibility to MFDs are considered. The modern concept of “disease ontogeny” (somatic mosaicism, loss of heterozygosity, paradominant inheritance, epigenetic variability) is discussed. It is demonstrated that the evolutionary and ontogenetic approaches to analysis of genimuc and other “-omic” data are essential for understanding the biology of diseases.     

楸树大小孢子发生与雌雄配子体发育的研究

运用石蜡切片法和整体透明法对楸树(Catalpa bungei C.A.Meyer)大、小孢子发生及雌、雄配子体发育过程进行了研究.结果表明:楸树可育雄蕊2枚,花药4室,药壁发育属双子叶型,腺质异型绒毡层.小孢子母细胞减数分裂为同时型,四分体后小孢子不分离形成正四面体型四合花粉,偶有左右对称型和十字交叉型.成熟花粉为二细胞型,无萌发孔.子房上位,2室,中轴胎座,胚珠多数,倒生,单珠被,薄珠心,具珠被绒毡层.单孢原直接发育为大孢子母细胞,四分体线形排列,合点端大孢子发育为功能大孢子,胚囊发育为蓼型.雄蕊发育早于雌蕊,花开后雌、雄蕊趋于同熟.研究认为:虽然楸树雌、雄蕊发育过程中均存在一定比例的败育,但其花而不实"并非雄性或雌性不育所致.推测与其授粉受精和胚后发育有关.