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Experimental and Cytological Studies on Plant Species

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Experimental and Cytological Studies on Plant Species Biologiske Skrifter udgivet af Det Kongelige Danske Videnskabernes Selskab Bind 11, nr. 5 Biol. Skr. Dan. Vid. Selsk. 11, no. 5 (1962) EXPERIMENTAL AND CYTOLOGICAL STUDIES ON PLANT SPECIES VII. GERANIUM SANGUINEUM BY TYGE W. BOCHER and MARTIN G. LEWIS D et Kongelige Danske Videnskabernes Selskab udgiver følgende pub­ likationsrækker : The Royal Danisii Academy of Sciences and L etters issnes the fol- lowing series of publications: Bibliographical Abbreviation Oversigt over Selskabets Virksomhed (8°) Overs. Dan. Vid. Selsk. (Annual in Danish) Historisk-filosofiske Meddelelser (8°) Hist. Filos. Medd. Dan. Vid. Selsk. Historisk-filosofiske Skrifter (4°) Hist. Filos. Skr. Dan. Vid. Selsk. (History, Philology, Philosophy, Archeology, Art History) Matematisk-fysiske Meddelelser (8°) Mat. Fys. Medd. Dan. Vid. Selsk. Matematisk-fysiske Skrifter (4°) Mat. Fys. Skr. Dan. Vid. Selsk. (Mathematics, Physics, Chemistry, Astronomy, Geology) Biologiske Meddelelser (8°) Biol. Medd. Dan. Vid. Selsk. Biologiske Skrifter (4°) Biol. Skr. Dan. Vid. Selsk. (Botany, Zoology, General Biology) Selskabets sekretariat og postadresse: Dantes Plads 5, København V. The address of the secretariate of the Academy is: Det Kongelige Danske Videnskabernes Selskab, Dantes Plads 5, Kbbenhavn V, Denmark. Selskabets kommissionær: E jn a r Mu n k sg a a r d ’s Forlag, Nørregade 6, København K. The publications are sold by the agent of the Academy: E jn a r M u n k sg a a r d , Publishers, 6 Nbrregade, Kobenhavn K, Denmark. Biologiske Skrifter udgivet af Det Kongelige Danske Videnskabernes Selskab Bind 11, nr. 5 Biol. Skr. Dan. Vid. Selsk. 11, no. 5 (1962) EXPERIMENTAL AND CYTOLOGICAL STUDIES ON PLANT SPECIES VII. GERANIUM SANGUINEUM BY TYGE W. BÜCHER and MARTIN C. LEWIS København 1962 i kommission hos Ejnar Munksgaard Synopsis The experimental material comprises 36 strains of Geranium sanguineum from various localities in Europe, grown under uniform conditions. All strains have the same chromosome number i.e. 2n = 84. Variation patterns investigated: (a) Leaf morphology. A cline in leaflobe width is found from E. to \V. Europe. A striking environmental modification of leaf shape is shown, (b) Leaf size. A cline from open to shaded habitats is found in the Danish material, (c) Petal moiphology. Variation in petal shape shows a degree of correlation with leaf shape, (d) Height and habit. Low coastal races and taller continental races are discernible. Three habit-types are found i. e. prostate, cushion and erect, (e) Flowering and vegetative phenology shows considerable variation which can be correlated with the climates of the native areas, (f) Hairiness. In Danish material there is a relationship between degree of hairiness and openness of the original habitat, (h) Flower colour. Although there is considerable variation, this is randomly distributed. Within this experimental material it is possible to recognize 2 race groups (I) Coastal race group (II) Woodsteppe race group. However these 2 groups only re­ present the ends of a more or less continuous clinal system. The taxonomic status of var. prostratum and var. lancastriense are considered. The significance of the variation in leaf morphology is discussed at some length. For several reasons it is believed that the leaf shape is of definite selective value: (I) The same distribution pattern of leaf types is found in several other species. (II) Variation in leaf shape is concurrent with ecotypic differentiation. (Ill) Narrow leaf lobing, such as is found in continental strains, is a well-known xeromorphic character, considered to be of selective value in dry climates. (IV) Environmental modification of the leaf parallels genotypically controlled adaption. PRINTED IN DENMARK BIANCO LUNOS BOGTRYKKERI A/S Introduction his paper is designed as another in the series of experimental studies undertaken Tby the senior author in cooperation with others. The purpose of the series is to supplement our taxonomic knowledge with information on the genetic-biological structure of the plant species. This involves studying genetic variability of the species, using information obtained from a study of living plants growing in cultivation. Inter-race patterns investigated include those involving the morphology, cytology and phenology of the species. Apart from the desirability of recording this variation, studies of this kind can be of value in revealing recent evolutionary trends within a species. In Geranium sanguineum, a striking variation in leaf shape was noted between strains from various parts of Europe. A more detailed study of this aspect was thought to be important, considering the taxonomic significance of leaf shape in the genus. Here, as in other genera (e.g. Ranunculus) leaf shape often provides the most dist­ inctive morphological character separating species. Variation in leaf morphology has proved useful in understanding evolutionary mechanisms in certain other genera. In this connection several investigations can be mentioned. W oodson (1947) was able to make a detailed analysis of introgression in the Asclepias tuberosa complex using leaf shape characters. Zimmermann’s work (1959) on the Pulsatilla aggregate illustrates how leaf characters can help in eluci­ dating recent history and modes of spéciation within a genus. In Taraxacum and Alchemilla it is possible to distinguish apomictic micro-species using small differences in leaf outline. Petersen’s examination of ecological leaf differentiation within the species Anthriscus sylvestris (1915, 1922), Potentilla erect a (1926) and Pimpinella saxifraga (1921) is particularly pertinent to the present paper and will be mentioned later in more detail. Leaf size has also proved useful in understanding the differentiation in tetra- ploid Veronica officinalis (B ôcher 1944). Genetically-determined differences in leaf area within this species form both a topocline from oceanic to continental regions and an ecocline from open to wooded localities. Other work on leaf morphology has centred mainly around morphogenetic problems. The striking changes in leaf morphology incident on submersion in water have received much attention. There is also a large volume of work on leaf xeromor- l* 4 Nr. 5 phism. Some plants, when grown under dry conditions will develop xeromorphic characters. In many plants, the upper leaves are more xeromorphic than those below. There has been much discussion about the adaptiveness of these structural changes to a water deficit (Maximov 1929; Shields 1950). However, only a few workers (e.g. Tukesson 1922, 1925; Fisher 1960) have investigated genetic variation in leaf shape within a species in terms of the selective response of the species to differences in environment. The junior author intends to study this aspect more fully using Geranium sanguineum and other species. In Veronica officinalis, leaf size forms “the backbone of the variational structure of the species” (B ocher 1944). In Geranium sanguineum, leaf shape fulfils the same role; it is central to an understanding of the organization within this species. In this paper other interracial patterns have been considered against this background. The characters investigated include petal morphology, height and habit, flower colour, flowering and vegetative periodicity, and hairiness. The senior author acknowledges with gratitude financial aid given by the Carls­ berg Foundation for the collection of material in various parts of Europe. The junior author expresses his thanks to N.A.T.O. Studentship Schemes for financial support during the period of this work. Material and Methods The experimental material comprises 36 strains of Geranium sanguineum from various localities in Europe (see Table I), grown under uniform conditions in the Experimental Garden of Copenhagen University. Field nos. 647-3195 have been in cultivation since 1953 - 3400-3533 - - - - - 1954 - 3796-4001 - - - - - 1955 - 4145-5111 - - - - - 1959 B 1-B 6 - - - - - 1959 The plants were grown from seed with the exception of strains 10, 14, 23, & 24 which were transplants from nature. Except in 4 numbers, 5-15 plants per strain were available. The leaf samples were pressed in 1960; 5 leaves were taken from each plant at different levels between the stem apex and base. The flower samples were harvested at the same time. Pressings of entire shoots were made in 1961; these were later used in studies on hairiness and leaf size. Root-tips from pot-grown plants were fixed in Miintzing’s modification of Navashin-Karpechenko and embedded in paraffin wax; these were later sectioned and stained in Feulgen’s Fuchsin. Root tips from strains 23, 24, and 32 were fixed in 3:1 Alcohol-Acetic after pretreatment in saturated aqueous solution of Monobromo- naphthalene; these were later stained in Feulgen’s Fuchsin and squashed. Nr. 5 5 T a b le 1. Country Field No. Locality Ecology Collector of origin No. Denmark 1 647 Dybdal, nr. Aalborg, N. Jutland ............... T. W. B. Steep grassy 2 648 Fosdal, nr. Fjerresslev, N. J u tla n d ........... — 3 649 Vilsund, N. Jutland ...................................... — 4 650 Boserup, nr. Roskilde, Zeeland..................... Wood edge — — 5 657 Skredbjærg, N. Zeeland................................ 1 Steep grassy 6 2073 ................................ slope — 7 2558 Dybesø, nr. Rørvig, N. Zeeland................... Stony rais. beach — Dry grassland K. Larsen 8 2563 Hirtsholmene, Island off N.E. Jutland. i 1 at the sea 9 2940 Bulbjærg, N. Ju tlan d ...................................... Fixed dunes T. W. B.
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