Embryology of Angiosperms

Thirty-four years have elapsed since the publication of the late Professor P. Maheshwari's text, An Introduction to the Embryology of Angiosperms, a work which for many years served as an invaluable guide for students and a rich source book for research workerso Various texts dealing with sections of the braad spectrum oftopics encompassed by Maheshwari in his book have appeared in the interim, but a compendious modem work dealing with the whole field has been lacking. This present volume splendidly meets the need, and it is altogether fitting that Professor B. M. lohri, long an associate and close colleague of Professor Maheshwari and himself a prolific contributor to the subject, should have undertaken the task of editing it. When Maheshwari wrote, it was stiIl feasible for one author to handIe the subject, but today even someone with his fine bread th of vision and depth of understanding could not, alone, do it justice. So the effort has to be a collaborative one; and Professor lohri's achievement has been to bring together a team of authoritative collaborators, assign them their responsibilities, and put them to work to produce a text as integrated in its treatment as the diversity of the subject would allow. The product vividly illustrates the advances that have been made in the study of angiosperm reproductive systems in the last 30 years, and the book is surely destined to become the new standard for student and researcher alike.

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1 Embryology: Then and Now.- 1.1 Introduction.- 1.2 Techniques to Study Embryology.- 1.3 Embryo Sacs in Anthers.- 1.3.1 Leptomeria.- 1.3.2 Calotis.- 1.3.3 Hyacinthus.- 1.4 Fritillaria Type of Embryo Sac.- 1.5 Unusual Embryological Features in Loranthaceae.- 1.5.1 Ovary-Ovule Complex.- 1.5.2 Embryo Sac.- 1.5.3 Endosperm.- 1.5.4 Embryo.- 1.5.5 Reproductive Calendar in Struthanthus vulgaris.- 1.6 Nutrition of the Ovule and Seed.- 1.6.1 Vascular Supply.- 1.6.2 Haustoria.- 1.6.3 Transfer Cells.- 1.7 Role of Persistent Pollen Tubes.- 1.8 Embryos of Doubtful Origin.- 1.8.1 Antipodal Embryos.- 1.8.2 Endospermal Embryos.- 1.9 Gymnospermous Characters in Angiosperms.- 1.9.1 Tambourissa.- 1.9.2 Butomopsis.- 1.9.3 Paeonia.- 1.10 Future Research.- References.- 2 The Microsporangium.- 2.1 Introduction.- 2.2 Morphology and Structure.- 2.3 Ontogeny.- 2.4 Wall Layers.- 2.4.1 Epidermis.- 2.4.2 Endothecium.- 2.4.3 Middle Layers.- 2.4.4 Tapetum.- 2.5 Ubisch Bodies/Orbicules.- 2.5.1 Origin.- 2.5.2 Relation to Exine Deposition: By-Product.- 2.6 Tapetal Membrane: Structure, Origin, and Significance.- 2.7 Pollenkitt and Tryphine.- 2.8 Sporogenous Tissue: Ultrastructure.- 2.8.1 Cytoplasmic Membranes and Ribosome Population.- 2.8.2 Nucleus.- 2.8.3 Nucleolar Cycle.- 2.8.4 Cytoplasmic Organelles During Meiosis.- 2.8.5 Plasmodesmata and Cytoplasmic Connections.- 2.9 Initiation and Control of Meiosis.- 2.9.1 Duration of Meiosis.- 2.9.2 Synthesis of Callose: Deposition and Significance.- 2.9.3 Cytokinesis.- 2.9.4 Cellulosic Wall of Microspore Mother Cell.- 2.10 Some Unusual Features.- 2.11 Conclusions.- References.- 3 The Ovule.- 3.1 Historical.- 3.2 Ovular Morphology.- 3.3 Ovule Initiation.- 3.4 Nucellus.- 3.5 Megasporogenesis.- 3.6 Integuments.- 3.7 Types of Ovules.- 3.8 Vascular Supply of the Ovule.- 3.9 Special Structures.- 3.10 Ovule Reduction.- 3.11 Concluding Remarks.- References.- 4 The Female Gametophyte.- 4.1 Introduction.- 4.2 Types of Megagametophyte Development.- 4.3 Development of the Megagametophyte.- 4.3.1 The Megaspore.- 4.3.2 The Coenocytic Megagametophyte.- 4.4 The Relation with the Nucellar Tissue.- 4.5 Organization of the Embryo Sac.- 4.5.1 The Synergids.- 4.5.2 The Egg Cell.- 4.5.3 The Central Cell.- 4.5.4 The Antipodal Cells.- 4.6 Female Gametophyte Development.- References.- 5 The Pollen Grain.- 5.1 Introduction.- 5.2 Pollen Structure and Cytochemistry.- 5.2.1 Pollen Shape.- 5.2.2 Pollen Cytology.- 5.2.3 Pollen-wall Structure.- 5.2.4 The Pollen Tube.- 5.2.5 Structural Adaptations of Pollen for Dispersal.- 5.3 Formation of Pollen.- 5.3.1 Cytology of Development.- 5.3.2 The Primexine.- 5.3.3 Establishment of Apertures.- 5.3.4 Exine Differentiation and Maturation.- 5.3.5 Intine Synthesis and Deposition.- 5.3.6 Mechanism of Cohesion in Composite Pollen.- 5.3.7 Pollen Sterility.- 5.4 Pollen Germination and Pistil Interactions.- 5.4.1 Pollen Quality.- 5.4.2 Pollen Germination and Tube Development.- 5.4.3 Diagnostic Landmarks of Pollination.- 5.4.4 Role of Pollen in Fertilization and Seed-setting.- 5.5 Pollen-wall Proteins and Allergens.- 5.6 Conclusions and Future Developments.- References.- 6 Fertilization.- 6.1 Introduction.- 6.2 Stigma and Style.- 6.2.1 Function and Structure of the Stigma.- 6.2.2 The Nature of the Stigma Covering.- 6.2.3 Function and Structure of the Style.- 6.3 Pollen Germination.- 6.3.1 Sticking and Hydration.- 6.3.2 Pollen Tube Formation and Growth.- 6.3.3 The Sperm Cell.- 6.4 Pollen-Pistil Interaction.- 6.4.1 Pollen Recognition, Acceptance, and Rejection.- 6.4.2 Pollen Incompatibility.- 6.4.3 Entry of Pollen Tube into Stigma and Style.- 6.4.4 Pollination Effects.- 6.5 Entrance and Discharge in Embryo Sac.- 6.5.1 Course of the Pollen Tube.- 6.5.2 Entry into the Embryo Sac.- 6.5.3 Growth Through the Filiform Apparatus.- 6.5.4 Entry into the Synergid.- 6.5.5 Transfer of Tube Content.- 6.6 Fusion of Gametes.- 6.6.1 Fusion of Nuclei.- 6.7 The Progamic Phase and Fertilization.- References.- 7 The Endosperm.- 7.1 Introduction.- 7.2 Nuclear Endosperm.- 7.2.1 Wall Formation.- 7.2.2 Cucurbitaceae.- 7.2.3 Leguminosae.- 7.2.4 Palmae.- 7.2.5 Proteaceae.- 7.2.6 Histochemistry and Ultrastructure.- 7.3 Cellular Endosperm.- 7.3.1 Acanthaceae.- 7.3.2 Cyrillaceae.- 7.3.3 Gesneriaceae.- 7.3.4 Icacinaceae.- 7.3.5 Santalaceae.- 7.3.6 Loasaceae.- 7.3.7 Loranthaceae.- 7.3.8 Scrophulariaceae.- 7.3.9 Histochemistry and Ultrastructure.- 7.4 Helobial Endosperm.- 7.4.1 Salient Features in Monocotyledons.- 7.4.2 Histochemistry and Ultrastructure.- 7.5 Ruminate Endosperm.- 7.6 Central Cell.- 7.6.1 Central Cell Cytoplasm.- 7.6.2 Central Cell Nucleus.- 7.6.3 Covering of the Central Cell.- 7.7 Wall Formation in Endosperm.- 7.8 Cytology of the Endosperm.- 7.9 Reserve Materials in Developing Endosperm.- 7.9.1 Protein Bodies.- 7.9.2 Starch.- 7.10 Embryo-Endosperm Relationship.- 7.11 Incorporation of Nucellus and Integument in Endosperm Formation.- 7.12 Conclusions and Prospects.- References.- 8 The Embryo.- 8.1 Historical.- 8.2 Zygote.- 8.2.1 Structure and Composition.- 8.2.2 Size Adjustments.- 8.2.3 Polarity.- 8.3 Early Embryogenesis.- 8.3.1 Cell Patterns.- 8.3.2 Tetrad, Quadrant, and Octant Proembryos.- 8.3.3 Stages Leading to Mature Embryo.- 8.4 Classification Based on Early Development of the Embryo.- 8.4.1 Systems Suggested by Schnarf and Johansen.- 8.4.2 System Suggested by Souèges.- 8.4.3 Other Systems of Classification.- 8.5 Differentiation in Embryo.- 8.6 Dicot and Monocot Embryo.- 8.7 The Grass Embryo.- 8.8 The Embryo in Palms.- 8.9 Ultrastructural and Cytochemical Aspects.- 8.10 Suspensor: Structure and Function.- 8.11 Deviations from Usual Development.- 8.11.1 Embryogeny in Paeonia.- 8.11.2 Embryos Devoid of Organs.- 8.11.3 Chimeral Embryos.- 8.12 Concluding Remarks.- References.- 9 Polyembryony.- 9.1 Introduction.- 9.2 Classification.- 9.2.1 Simple Polyembryony.- 9.2.2 Multiple Polyembryony.- 9.3 Nucellar Polyembryony.- 9.3.1 Rutaceae.- 9.3.2 Anacardiaceae.- 9.3.3 Myrtaceae.- 9.3.4 Cactaceae.- 9.3.5 Orchidaceae.- 9.4 Integumentary Polyembryony.- 9.4.1 Endothelial Polyembryony.- 9.5 Zygotic and Suspensor Polyembryony.- 9.5.1 Zygotic Polyembryony.- 9.5.2 Suspensor Polyembryony.- 9.6 Synergid Polyembryony.- 9.6.1 Twins and Triplets.- 9.7 Polyembryony in Hybrids.- 9.8 Causes of Polyembryony.- 9.9 Induction of Polyembryony.- 9.10 Utilization of Plural Embryos.- References.- 10 Gametophytic Apomixis.- 10.1 Introduction.- 10.1.1 Definitions.- 10.1.2 Synonyms.- 10.1.3 Remarks on Terminology.- 10.2 Embryology of Gametophytic Apomicts.- 10.2.1 Development of Unreduced Embryo Sacs.- 10.2.2 Formation of Endosperm and Embryo.- 10.2.3 Male Meiosis and the Male Gametophyte.- 10.3 Apomixis and Sexuality.- 10.3.1 Sexual Potentials in Gametophytic Apomicts.- 10.3.2 Possibilities of Influencing the Degree of Apomixis.- 10.4 Causes and Consequences of Apomixis.- 10.4.1 Heterozygosity and Variability.- 10.4.2 Causes of Gametophytic Apomixis; Components of Apomixis in Sexual Plants.- 10.4.3 Apomixis and Polyploidy.- 10.4.4 Agamic Species Complexes and Their Evolution.- 10.5 Genetics of Gametophytic Apomixis.- 10.5.1 Dominant or Recessive Inheritance?.- 10.5.2 Some Critical Remarks.- 10.5.3 Some Results, Conclusions, and Prospects.- 10.6 Apomixis and Breeding Conclusions.- References.- 11 Role of Polyploidy in Reproductive Organs and Tissues.- 11.1 Introduction.- 11.2 Methods of Polyploidization and Ploidy Determination.- 11.2.1 Acytokinetic Mitosis (AM).- 11.2.2 Spindle Fusion (SF).- 11.2.3 Restitutional Mitosis (RM).- 11.2.4 Endomitosis (Em).- 11.2.5 Chromosome Endoreduplication (Er).- 11.2.6 Ploidy Determination.- 11.3 Anther.- 11.3.1 Anther Wall and Connective.- 11.3.2 Tapetum.- 11.3.3 Pollen Grains.- 11.4 Ovule.- 11.4.1 Nucellus.- 11.4.2 Integument(s) and Endothelium.- 11.5 Embryo Sac.- 11.5.1 Egg Cell.- 11.5.2 Synergids.- 11.5.3 Proendospermic Cell.- 11.5.4 Antipodal Cells.- 11.6 Endosperm.- 11.6.1 Origin and Types of Endosperm.- 11.6.2 Polyploidy in the Endosperm.- 11.7 Embryo.- 11.7.1 Embryo Proper.- 11.7.2 Suspensor.- 11.7.3 Nonzygotic Embryos.- 11.8 Physiological Role of Polyploidy.- 11.9 Future Research.- References.- 12 The Seed: Structure.- 12.1 Introduction.- 12.1.1 Importance of Seeds.- 12.1.2 Historical Survey.- 12.2 Structure of Seeds.- 12.2.1 General Morphology.- 12.2.2 Vascular Supply of the Seed.- 12.2.3 Seed Types.- 12.2.4 Development and Structure of Seed-coats.- 12.2.5 Differentiation.- 12.2.6 The Seed-coat Surface.- 12.2.7 Stomata in Seed-coats.- 12.3 Special Structures, Adaptations, and Functions.- 12.3.1 Sarcotestal Seeds.- 12.3.2 Arillate Seeds.- 12.3.3 Jaculator.- 12.3.4 Winged Seeds.- 12.3.5 Haired Seeds.- 12.3.6 Dust and Balloon Seeds.- 12.3.7 Hydrochoric Seeds.- 12.3.8 Mucilage Seeds.- 12.3.9 Active Dispersal of Seeds (Autochoric Seeds).- 12.3.10 Color of Seeds.- 12.4 Adaptations to Germination.- 12.4.1 Opercula.- 12.4.2 Imbibition Lids.- 12.4.3 Epistase and Hypostase.- 12.5 Rumination and Labyrinth Seeds.- 12.5.1 Labyrinth Seeds.- 12.6 Storage Tissue.- 12.7 Concluding Remarks.- References.- 13 The Seed: Germination.- 13.1 Introduction.- 13.2 General Considerations.- 13.2.1 Internal Morphology.- 13.2.2 Composition of Seeds and Reserve Foods.- 13.3 Germination: Structural, Compositional, and Physiological Changes.- 13.3.1 Dicotyledons.- 13.3.2 Monocotyledons.- 13.4 Storage and Aging of Seeds: Structure and Physiology.- References.- 14 Embryology and Taxonomy.- 14.1 Introduction.- 14.2 Embryological Features of Taxonomic Importance.- 14.3 Applications of Embryology to Problems of Family Relationships.- 14.3.1 Podostemaceae.- 14.3.2 Basellaceae.- 14.3.3 Stachyuraceae.- 14.3.4 Frankeniaceae.- 14.3.5 Tropaeolaceae.- 14.3.6 Salvadoraceae.- 14.3.7 Centrolepidaceae.- 14.4 Applications of Embryology to Problems of Taxonomic Structure Within Families.- 14.4.1 Aizoaceae.- 14.4.2 Vacciniaceae.- 14.4.3 Zannichellia.- 14.5 Applications of Embryology to Problems of Generic Affinity and Assignment.- 14.5.1 Ploiarium.- 14.5.2 Filicium.- 14.5.3 Nyctanthes-Jasminum.- 14.5.4 Ophiopogon.- 14.5.5 Kyllinga.- 14.5.6 Burmannia pusilla.- 14.6 Contributions of Embryology to Taxonomic Problems Still Unsolved.- 14.6.1 Erythroxylaceae-Linaceae.- 14.6.2 Theligonaceae.- 14.6.3 Blandfordia.- 14.6.4 Philydraceae.- 14.6.5 Eriocaulaceae.- 14.7 Confirmation or Alteration of Earlier Conclusions.- 14.7.1 Cactaceae.- 14.7.2 Garryaceae.- 14.7.3 Orobanchaceae.- 14.7.4 Pentaphragma.- 14.8 Embryology in Relation to Taxonomy - the Prospect.- 14.8.1 Cornus.- 14.8.2 Glycine and Phaseolus.- 14.8.3 Eustachys petraea and E. glauca.- References.- 15 Homologies and Phylogeny.- 15.1 Introduction.- 15.1.1 Sexuality and Overall Phylogeny.- 15.2 Microsporangia.- 15.2.1 Microsporangium Wall.- 15.2.2 Microsporangium Content.- 15.2.3 Microsporangium Number and Position.- 15.3 Megasporangia.- 15.3.1 General Structure.- 15.4 Ontogeny of Nucellus.- 15.4.1 Tapetum/Endothelium.- 15.4.2 Megasporangium Content.- 15.4.3 A Proposed Macroevolution of Meiosporic Structures.- 15.5 Male Gametophyte.- 15.5.1 Extrinsic to Angiosperms.- 15.5.2 Intrinsic to Angiosperms.- 15.6 Female Gametophyte.- 15.6.1 Female Sexual Structures in Chlorophyta Extrinsic to Angiosperms.- 15.6.2 Embryo Sac.- 15.6.3 Diversity in Embryo Sacs.- 15.6.4 Ontogeny of Embryo Sac as Compared to Female Gametophyte of Taxus.- 15.7 Pollen Reception by Female Organ.- 15.7.1 Fertilization.- 15.7.2 Double Fertilization, Extrinsic to Angiosperms.- 15.7.3 Intrinsic to Angiosperms.- 15.7.4 Cytology of Zygote Formation.- 15.8 Embryogenesis.- 15.8.1 Albuminogenesis (or Endosperm Formation) in Angiosperms.- 15.8.2 Embryo.- 15.9 Seed.- 15.9.1 Time Relations in the Development of Seed.- 15.9.2 Diversity in Seed Structure.- 15.10 Concluding Remarks.- 15.10.1 Questions Raised by Natural Diversities.- 15.10.2 Questions Raised by Suggested Experiments.- References.- 16 Experimental Embryology.- 16.1 Introduction.- 16.2 The Problems: Role of Experimental Embryology.- 16.3 Techniques of Cell, Tissue, and Organ Culture.- 16.4 Floral Organs and Buds.- 16.4.1 In Vivo Studies.- 16.4.2 In Vitro Studies.- 16.5 Anther and Pollen Culture.- 16.5.1 Cultural Procedure and Nutritional Requirements.- 16.5.2 Anther Culture.- 16.5.3 Pollen Culture.- 16.5.4 The Pollen Plants.- 16.5.5 Use of Haploids.- 16.6 Ovary, Ovule, and Nucellus Culture.- 16.6.1 Ovary Culture.- 16.6.2 Ovule Culture.- 16.6.3 Nucellus Culture.- 16.7 Control of Fertilization.- 16.7.1 Intra-ovarian Pollination and Fertilization.- 16.7.2 In Vitro Pollination of Ovules and Fertilization.- 16.7.3 Inter- and Intrageneric Hybrids.- 16.8 Endosperm Culture.- 16.8.1 Immature Endosperm.- 16.8.2 Mature Endosperm.- 16.8.3 Embryo-Endosperm Relationship.- 16.8.4 Organogenesis in Endosperm Cultures.- 16.9 Embryo Culture.- 16.10 Polyembryony.- 16.11 Control of Embryogenesis.- 16.11.1 Loss of Embryogenic Potency.- 16.11.2 Ontogeny of Embryos.- 16.12 Protoplast Culture.- 16.12.1 Embryogenesis and Regeneration.- 16.12.2 Protoplast Fusion and Somatic Hybridization.- 16.12.3 "Pomatoes" and "Topatoes".- 16.13 Concluding Remarks.- References.- Plant Index.