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The Morphogenesis of the Inflorescence, Flower and Fruit of Pyrus Nivalis Jacquin Var. Orientalis Terpó

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The Morphogenesis of the Inflorescence, Flower and Fruit of Pyrus Nivalis Jacquin Var. Orientalis Terpó 582.734.3:581.145:581.46:581.47 MEDEDELINGEN LANDBOUWHOGESCHOOL WAGENINGEN • NEDERLAND • 70-5 (1970) THE MORPHOGENESIS OF THE INFLORESCENCE, FLOWER AND FRUIT OF PYRUS NIVALIS JACQUIN VAR. ORIENTALIS TERPÓ G. STARITSKY Afdeling Tropische Plantenteelt, Landbouwhogeschool, Wageningen, Nederland. (Received24-IX-1969 ) H.VEENMA N & ZONEN N.V.-WAGENINGEN-1970 Mededelingen Landbouwhogeschool Wageningen 70-5 (1970) (Communications Agricultural University) isals o published as a thesis CONTENTS 1. INTRODUCTION 1 2. MATERIAL AND METHODS 2 2.1. Material 2 2.2. Sampling,measurement s and observation 3 2.3. Microtechnique 3 3. GENERAL SECTION 5 3.1. Classification 5 3.2. Morphology 6 3.2.1. Structure ofth etre e 6 3.2.2. Inflorescence structure and type 8 3.2.3. Initiation and development ofth einflorescenc e 10 3.2.4. The flower 14 3.2.5. Thefruit 17 4. THEDEVELOPMEN T OFTH EINFLORESCENC E ANDFLOWE R 19 4.1. Introductory 19 4.2. Theinitiatio nan d development ofth emixe dbu d 19 4.2.1. Surveyo fth edevelopmen t 19 4.2.2. Thevegetativ ephas e 26 4.2.3. Commencement ofth egenerativ ephas e 27 4.2.4. Autumn development ofth einflorescenc e 30 4.2.5. Winter development ofth einflorescenc e 31 4.2.6. Springdevelopmen t ofth einflorescenc e 33 4.3. Correlation betweenflower characteristic s and positioni nth einflorescenc e . 34 4.3.1. Correlation between ovarydiamete r and positioni nth einflorescenc e .... 34 4.3.2. Thenumbe r ofstamen spe r flower 36 4.3.3. Number ofstyle spe r flower 37 4.3.4. Number of styles in the flower in relation to the number of locules in the fruit 38 4.3.5. Number ofstyle san dlocule si nflowers o fdevelopin ginflorescence s 39 4.3.6. Theformatio n ofquadriloculat eflowers wit hfiv estyle s 40 4.4. Surveyo fth edevelopmen t ofth einflorescenc e and flower 42 5. GROWTH AND DEVELOPMENT OFTH EFRUI T 44 5.1. Introductory 44 5.1.1. Growth 44 5.1.2. Development 44 5.1.3. Morphogenesis 45 5.1.4. Histogenesis (sclereid clusters,intervenin gparenchym a cells) 46 5.2. Fruit growth 48 5.2.1. Increasei ndiamete r 48 5.2.2. Increasei nvolum e 52 5.2.3. Fresh weightan d dryweigh t 55 5.3. Morphogenesis 57 5.3.1. Pathenocarpy 57 5.3.2. Seedlessnessan ddimensio n 57 5.3.3. Seedlessness and shape 59 5.3.4. Seedlessnessan d coresiz e 60 5.3.5. Thecaus eo fflattening o fseedles sfruit s 61 5.3.6. Formula for thelength/diamete r ratio 65 5.3.7. Correlation betweenfrui t sizean d thepositio n inth einflorescenc e 66 5.3.8. Correlation betweenfrui t production and thepositio n inth einflorescenc e . 69 5.3.9. Number ofseed si nth efruit s 70 5.3.10. Numbero flocule si nth efruit s 71 5.3.11. Differences ingrowt h on different branches 72 5.4. Histogenesis 73 5.4.1. Number ofsclerei dcluster si nth erip epea r 73 5.4.2. Number ofsclereid spe rcluste r andestimate s ofnumbe r ofsclereid spe rpea r . 75 5.5. Surveyo fth efrui t development 78 SUMMARY 81 ACKNOWLEDGEMENTS 84 SAMENVATTING 85 REFERENCES 88 1. INTRODUCTION In his articles on the physiology of fruit development, NITSCH (1953, 1965) rightly observes that fruit growth does not begin at pollination of the flower but at an earlier stage during initiation of the floral primordia. He distinguishes two important periods ofgrowt h inth e life-history of the fruit, viz.: 1. the period of pre-bloom development when growth mainly occurs through celldivisio n; 2. the post-fertilisation period when cell enlargement is the most important factor. These two periods are separated by the critical phases of anthesis and pollina­ tion. As far as the first period is concerned, flower initiation has been studied in extenso (HILLMAN, 1962; LANG, 1965; SALISBURY, 1963). Many research workers believe that after initiation of the floral primordia the differentiation and development of the various parts of the flower is a more or less automatic process following a well-marked pattern. Owing to this misconception the study of these subjects has been greatly neglected. The second period is one of substancial growth of the young fruit, which explainswh y this is often the only period considered in fruit-growth studies. As a result certain important facts providing a better understanding of the nature of fruit growth may wellhav e been overlooked. The object of the present study of the growth of the fruits of Pyrus nivalis Jacquin, a wild species of pear distributed over Central Europe, the Danube countries, the Balkan Peninsula and Asia Minor, is to place proper emphasis on the morphogenesis of both the flower and fruit so as to provide the fullest possible information on this subject. The literature contains no references to the fruit development of wild pear species, the only comparable literature available relating to the fruit develop­ ment of cultivars. Furthermore, compared with other fruit-trees, surprisingly little research work has been done on pears. This may be illustrated byth ebib ­ liography by SCARAMUZZI (1953) who studied the bud development of the plum,peach , cherry, apple and pear. Out of a total of 237 bibliographical ref­ erences, 50 relate to the apple, two to apple and pear together, but only five to the pear alone. Out of a total of 370 references in NITSCH (1953), 30 relate to the apple, 6t o apple and pear, and only three to thepea r alone. As regards the study of Pyrus nivalis,th e following topics will be discussed in succession: 1. the initiation of theflower primordi a; 2. the differentiation ofth e various parts of the flower; 3. the development ofth e ovary prior to flowering; 4. anthesis; 5. the development of the ovary and any other parts of the flower to the ripe fruit. Meded. Landbouwhogeschool Wageningen 70-5 (1970) 1 2. MATERIAL AND METHODS 2.1. MATERIAL A very old pear-tree may be seen on the rampart of the Leiden Botanical Gardens. In a plan drawn by H. WITTE in 1868 the tree shown on this site is termed Pyrus sinaica (VEENDORP and BAAS BECKING, 1938), and this name oc­ curs earlier in catalogues of the Botanical Gardens, i.e. in BRUGMANS' catalogue (1819) under the author's name Desf., in REINWARDT'S catalogue (1831), and in a manuscript catalogue dated 1861 under the author's name THOIN. The tree was probably planted in the newly acquired part after the fifth extension in 1816. Grafts from the tree were brought to the Arboretum 'De Dreijen' at Wage­ ningen in 1935.Th e tree that developed from a graft was determined by HENSEN (1962),wit h the assistance of TERPÓ'S article (1960),a s Pyrus nivalisJacqui n var. orientalis Terpó. Besides this tree Wageningen has three young specimens of the same variety in the Belmonte Arboretum. These young trees, which were planted in 1957, were also determined by HENSEN (1962). One of the three, hereafter referred to as tree I, bears a striking resemblance to the Leiden tree. The two other young trees, II and III, are somewhat different (among other characters the flowers and fruits are larger). Moreover the vegetative growth of trees II and III is far greater than of tree I whose shoots, like those of the Leiden tree, only rarely make considerable extension growth. The vigorously growing trees were possi­ bly raised from seed, whereas the slowly growing tree I may come from a graft taken from thelarg e tree at Leiden. Material from thelarg etre ea t Leiden wasmeasure d and sampled from 1958t o 1963. The Wageningen material was used for research in 1964 and subsequent years. Close to the large tree on the rampart at Leiden is a smaller pear-tree named Pyrus nivalis. According to HENSEN'S (1962) determination it should be known as Pyrus austriaca Kern. (Syn. P. nivalis Jacq, f. austriaca (Kern.) Schneid.). In sizean d structure of itsfruit s it differs from the large tree. A few complementary studies have been devoted to other pear-tree species found in the Belmonte Arboretum. The names accompanying the trees in the Arboretum have been adopted in the present publication without modification. The species in question are: Pyrus amygdaliformis Vill., Pyrus austriaca Kern., Pyrus calleryanaDene , Pyrus elaeagrifoliaPall. , Pyrus domestica Med., Pyruspyrifolia (Burm.fil.) Nak . var. culta (Makino) Nak., Pyrus salicifoliaPall. , and Pyrus ussuriensisMaxim , var. ovoidea (Rehd.) Rehd.. 2 Meded. Landbouwhogeschool Wageningen 70-5 (1970) 2.2. SAMPLING, MEASUREMENTS ANDOBSERVATIO N In their study of the apple TUKEY and YOUNG (1942) found that it is much more difficult tosecur e samplings ofthi s fruit asunifor m aspossibl e than those of the cherry, forexample .Clos einspectio n ofth efruits , thebuds , inflorescences and flowers of Pyrus nivalis also reveals fairly great differences. Although a great many samples should be taken for this very reason, relatively little material was collected to avoid damaging theplan t andupsettin g the balance. This is particularly true of fruits whose small number in certain years ofth e study was due toa poo r fruit-set. Samples were therefore nottake n at random; instead the specimens selected were, as far as possible, representative of the development. After sampling, the buds, inflorescences, flowers and fruits were fixedi n 70 % alcohol, a formol - alcohol - acetic acid solution (FAA) in which half the acetic acid is sometimes replaced by propionic acid (FAPA), or in a chromic acid - acetic acid - formol solution of varying ratios of constituents (Crafl-VinSASS, 1961).
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