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Morphological Studies Op Diploid Aud Autotetraploid MORPHOLOGICAL STUDIES OP DIPLOID AUD AUTOTETRAPLOID PLARTS OP PHYSALIS PRUIUOSA L. Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By ROBERT DAVID HEURY, B.S., M.S. The Ohio State University 1958 Approved hy Department of Botany and Plant Pathology ACKN OWLEDGMEUT S The writer desires to take this opportunity to express his sincere thanks and appreciation to his adviser Dr, G, W. Blaydes for his guidance, advice, criticisms, and encouragement during the course of this investigation and preparation of the dissertation. Thanks are also extended to Dr. K. K. Pandey for his helpful suggestions concerning the research and to Mr. A. S. Heilman for the photographic work. ii TABLE OP COM'Ei'TTS Page IHTRODU CTI ON ...................................... 1 MATERIALS ARB M E T H O D S ........................ 4 GENERAL MORPHOLOGY AND G R O W T H ............... 8 Observations and Results ..................... 8 Discussion and Summary............ 21 PRE- ADD POST-PERTILIZATIOH MORPHOLOGY............. 27 Development of the Ovule and Emhryo Sac .... 27 Observations and Results ............. 27 Discussion and Summary ................... 33 Development of the Pollen ............ 37 Observations and Results ................. 37 Discussion and Summary ............... 42 Fertilization ........ ................. 43 Observations and Results • ••••...• 43 Discussion and Summary ............. 44 Endosperm ........ .............. 43 Observations and Results .......... 45 Discussion and Summary ................... 48 Embryogeny .... ............... ...... 49 Observations, Results and Discussion . 49 Summary 57 Early Seed Coat Development •••••••••• 58 iii Table of Contents - Continued. Page AUTOTETRAPLOID STERILITY ......................... 60 Observations and Results ............ 6l Discussion and Summary ............ 70 PAETHENOCARPY . ................ 74 SUMMARY ..................... 79 LITERATURE CITED ................................... 82 iv LIST OF TABLES Table Page 1 Size of leaf tissues and size and frequency of stomates in diploid and" tetraploid Ph.ysalis pruinosa ..... ......................... 11 2 Flower and fruit characteristics of diploid and tetraploid Physalis pruinosa- ........ 13 3 Relation of fruit size to number of seeds in diploid and tetraploid Physalis pruinosa . 18 4 Seed germination of diploid and tetraploid Physalis pruinosa ..................... 18 5 Rate of growth in height of diploid and tetra­ ploid Physalis pruinosa ..... .......... 20 6 Comparison between autotetraploid Physalis pruinosa and autotetraploid Physalis floridana . ........ 22 7 Comparative data on the embryo sac of diploid and tetraploid Physalis pruinosa...... 36 8 Fertilization and pre- and post-fertilization abortion in diploid and tetraploid Physalis pruinosa............................ 68 9 Size of mature fruits and associated structures in diploid and tetraploid Physalis pruinosa formed as a result of self-pollination in comparison to 0.25 per cent naphthaleneacetic acid being applied to the stigma............ 75 v LIST OP ILLUSTRATIONS (All illustrations are of Physalis pruinosa L e) Figure Page 1 Metaphase chromosomes in pollen mother cell of d i p l o i d ............................ 5 2 Metaphase chromosomes in pollen mother cell of tetraploid . .............. • ........... 5 3 Diploid and tetraploid plants • 10 4 Leaf cross-section of d i p l o i d .................... 12 5 Leaf cross-section of tetraploid .................. 12 6 Flowers of diploid and tetraploid ................ 14 7 Seeds of diploid and t e t r a p l o i d ........... 14 8 Pollen of diploid ........................ 15 9 Pollen of tetraploid .......... * ............. 15 10 Megaspore mother cell, diploid .................... 31 11 Dyad (megaspore), diploid ............. 31 12 Tetrad of megaspores, d i p l o i d .................... 31 13 Functional megaspore, diploid ..... .......... 31 14 2-nucleate embryo sac, diploid .......... • . • 32 15 4-nucleate embryo sac, diploid 32 16 8-nucleate embryo sac, diploid 32 17 Microspore mother cell, tetraploid .......... 39 18 Dyad (microspore), tetraploid .......... ..••• 39 19 Tetrad of microspore nuclei, tetraploid . • . 40 20 Tetrad of microspores, tetraploid ........ 40 21 2-nucleate pollen, tetraploid 41 22 Microcytes in microspore tetrad of tetraploid . 41 vi List of Illustrations - Continued Figure Page 23 2-celled endosperm, diploid ....... 47 24 Embryo with. 8 cells in 6 tiers, d i p l o i d ............. 55 25 Embryo with 8 cells in 8 tiers, diploid ....... 55 26 Embryo with 8 cells in 7 tiers, d i p l o i d ............ 55 27 Embryo with 5 cells in 4 tiers, d i p l o i d ............. 55 28 Embryo with 6 cells in 6 tiers, 1 in metaphase, -diploid ...... ................................ 55 29 Embryo with 8 cells in 6 tiers, a diploid variation . 56 30 Aberrant 6-celled embryo of diploid ......... .... 56 31 Embryo with 8 cells in 6 tiers, a tetraploid variation .............................. 56 32 Embryo with 8 cells in 7 tiers, a tetraploid v a r i a t i o n ................................. 56 33 Embryo with 8 cells in 6 tiers, a tetraploid v a r i a t i o n ........................... 56 34 Pollen tubes of tetraploid at base of style of tetraploid . ......... ............ 67 35 Pollen tubes of diploid at base of style of diploid . 67 36 Fruits of diploid as a result of self-pollination and of hormone placed on the stigma ......... 76 37 Fruits of tetraploid as a result of self-pollination and of hormone placed on the stigma •••••.• 76 vii INTRODUCTION Physalis is one of the genera of the Solanaceae, a family which contains some economically important crops including tomatoes, potatoes and eggplant# Although not generally considered of great economic importance, some species, including P. pruinosa# are culti­ vated for their fruits which are used in jams and relishes. The Chinese lantern, P. alkekengi, is often cultivated for its orna­ mental calyces when in fruit. Physalis is often found as a common weed in temperate and tropical regions of the United States. In this study reference will he made to some of the many morphological studies in the Solanaceae, hut apparently there has heen very little morphological work done in this genus. In 1896 it was monographed hy Rydherg and recently a cytological-genetic investigation was completed hy Menzel (l95l)» The megagametophyte of P . minima and P. peruviana as well as the emhryogeny of P. minima has heen studied hy Bhaduri (1935» 1936). Tognini (1 9 0) 0 described a few stages in the emhryogeny of P. edulis and Mascrd (1921) conducted some studies of the anther in some genera of the Solanaceae including Physalis. Much has heen written on the utilization of tetraploids and other polyploids in plant breeding and agriculture hy Eigsti (1957)» Eigsti and Dustin (l955)» Dermen (1940), Emsweller and Ruttle (1941)t Stebbins (1956) and others. As these workers often mention, the advantages such as larger ecxmomically important plant organs and use in breeding for introduction of qualitative and quantitative changes in hybrids, are often offset by such disadvan- tages as slower growth, pollen sterility, and, particularly, low seed set. A study of Physalis pruinosa was conducted to contribute to the morphological knowledge of the genus as well as of the family. For example, there has heen some question on the development of the mega garnetophyte in this family (Schnarf 1931> Bhaduri 1935) • As a result of an accumulation of morphological information from the various investigations of many workers, a better understanding of the systematic and phylogenetic relationships may be obtained. A knowledge of the morphology may help in the analysis and understand­ ing of problems frequently encountered in commercial crop production in such an economically important family. The results of studies conducted on the development of the ovule, embryo sac, pollen, endosperm, young embryo and seed coat of Physalis pruinosa L. are reported in this dissertation. In addition, due to the increased interest in polyploidy in agriculture, an investigation on the comparative morphology of diploid and colchicine-induced autotetraploid plants of Physalis pruinosa. including a study on the cause of low seed set in the latter, was thought to be a desirable addition to the morphology of polyploids that has accumulated and which in some cases has been inconsistent• A preliminary study made on parthenooarpic fruit develop­ ment in this species is also contained in this report. In some economically important fruits, including grape®, oranges and grape­ fruit among others, parthenocarpy has been important. If there is 3 an increase in the utilization of Physalis as a fruit crop, parthe- nocarpy may he desirable# There is an abundance of literature on the subjects studied in this investigation.' Obviously, it is impossible to cite all of them. In the introduction to the major divisions of this study (i.e., General Morphology, and Growth, Pre- and Post-fertiliza' tion Morphology, Autotetraploid Sterility, and Parthenocarpy) re­ ference is made to some relevant reviews and comprehensive works0 Along with the observations and discussion of the results of -this study are included citations to some of the other works that are relevant to the particular topic at hand. References cited are often limited to the Solanaceae since this is the family concerned in this study. MATERIALS AND METHODS Diploid and autotetraploid
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