<p>The Holocene spread of <i>Picea abies</i> in Scandinavia provides an excellent opportunity for detailed study of the dynamics of tree spread and population expansion. Early- and mid-Holocene macrofossil evidence for the presence of <i>Picea abies</i> in Scandinavia has questioned traditional interpretations of the timing and direction of its spread. This study aims to determine the pattern of the spread of <i>Picea abies</i> in Scandinavia from pollen and other data, to evaluate the significance of possible early outpost populations and to deduce possible factors that influenced the spread and population expansion of <i>Picea abies</i> in Scandinavia. </p><p>Palaeoecological investigations were carried out on the sediments of four small lakes in central Sweden to gain detailed insight into the dynamics of the spread. Holocene pollen diagrams with independent dating control were collected from Fennoscandia and adjacent areas to compare the timing of selected features of the <i>Picea abies</i> pollen curve. Computer models were used to test possible scenarios for the spread and <i>Picea abies</i> population expansion. </p><p><i>Picea abies</i> entered the Scandinavian peninsula from the east at different times and by different pathways. Early-Holocene outposts can be discerned in pollen records from northwest Russia, eastern and northeastern Finland for the time before 9000 cal. BP. Pollen records from Sweden and Norway indicate small <i>Picea abies</i> populations after 8000 cal. BP. The mid to late-Holocene spread, which superficially resembles a front-like pattern, may in fact represent a wave of expanding populations. Disturbance through fire and human activity did not significantly influence the pattern of the spread. Changing climate parameters, slow adaptation and gene flow through seeds and pollen have to be considered as possible explanations for the late spread of the tree. Population dynamics and propagule pressure are likely to be important factors that shaped the spread of <i>Picea abies</i>.</p>

<p>The reproductive organs of conifers and angiosperms differ in morphology in several fundamental respects. The conifer Norway spruce <i>(Picea abies)</i> form pollen and seed cones from separate meristems whereas angiosperms bear bipartite flowers with sepals and petals surrounding two inner whorls of stamens and carpels. Despite these differences in morphology this thesis present data to suggest that reproductive development in conifers and angiosperms is regulated by a similar molecular mechanism. This implies an evolutionary conservation of the major mechanism for reproductive development since the origin of seed plants. </p><p>Flower organ identity in angiosperms is determined by regulatory genes belonging to the MADS-box gene family of transcription factors. This thesis presents the cloning and characterisation of four novel MADS-box genes from Norway spruce<i>.</i> Three of these genes <i>DAL11</i>, <i>DAL12</i> and <i>DAL13 </i>are most closely related to angiosperm B function genes <i>i.e.</i> genes required for petal and stamen development. <i>DAL11</i>, <i>12</i> and <i>13</i> all are specifically active in developing pollen cones, with different temporal and spatial expression pattern. Functional analysis in transgenic Arabidopsis and yeast suggest that the reproductive aspect of the B-function is conserved between conifers and angiosperms. The results also suggest that the B-function in conifers is separated into one shoot identity and one organ identity determinant. </p><p>A fourth gene presented;<i> DAL10,</i> is specifically expressed in vegetative parts of pollen- and seed cones. Phylogenetically <i>DAL10</i> is not closely related to any of the known angiosperm clades, but rather forms a separate clade with other gymnosperm genes, suggesting a gymnosperm specific function. We suggest that the <i>DAL10</i> activity reflects a function in the determination of the reproductive shoot.</p>

The reproductive organs of conifers and angiosperms differ in morphology in several fundamental respects. The conifer Norway spruce (Picea abies) form pollen and seed cones from separate meristems whereas angiosperms bear bipartite flowers with sepals and petals surrounding two inner whorls of stamens and carpels. Despite these differences in morphology this thesis present data to suggest that reproductive development in conifers and angiosperms is regulated by a similar molecular mechanism. This implies an evolutionary conservation of the major mechanism for reproductive development since the origin of seed plants. Flower organ identity in angiosperms is determined by regulatory genes belonging to the MADS-box gene family of transcription factors. This thesis presents the cloning and characterisation of four novel MADS-box genes from Norway spruce. Three of these genes DAL11, DAL12 and DAL13 are most closely related to angiosperm B function genes i.e. genes required for petal and stamen development. DAL11, 12 and 13 all are specifically active in developing pollen cones, with different temporal and spatial expression pattern. Functional analysis in transgenic Arabidopsis and yeast suggest that the reproductive aspect of the B-function is conserved between conifers and angiosperms. The results also suggest that the B-function in conifers is separated into one shoot identity and one organ identity determinant. A fourth gene presented; DAL10, is specifically expressed in vegetative parts of pollen- and seed cones. Phylogenetically DAL10 is not closely related to any of the known angiosperm clades, but rather forms a separate clade with other gymnosperm genes, suggesting a gymnosperm specific function. We suggest that the DAL10 activity reflects a function in the determination of the reproductive shoot.

The Holocene spread of Picea abies in Scandinavia provides an excellent opportunity for detailed study of the dynamics of tree spread and population expansion. Early- and mid-Holocene macrofossil evidence for the presence of Picea abies in Scandinavia has questioned traditional interpretations of the timing and direction of its spread. This study aims to determine the pattern of the spread of Picea abies in Scandinavia from pollen and other data, to evaluate the significance of possible early outpost populations and to deduce possible factors that influenced the spread and population expansion of Picea abies in Scandinavia. Palaeoecological investigations were carried out on the sediments of four small lakes in central Sweden to gain detailed insight into the dynamics of the spread. Holocene pollen diagrams with independent dating control were collected from Fennoscandia and adjacent areas to compare the timing of selected features of the Picea abies pollen curve. Computer models were used to test possible scenarios for the spread and Picea abies population expansion. Picea abies entered the Scandinavian peninsula from the east at different times and by different pathways. Early-Holocene outposts can be discerned in pollen records from northwest Russia, eastern and northeastern Finland for the time before 9000 cal. BP. Pollen records from Sweden and Norway indicate small Picea abies populations after 8000 cal. BP. The mid to late-Holocene spread, which superficially resembles a front-like pattern, may in fact represent a wave of expanding populations. Disturbance through fire and human activity did not significantly influence the pattern of the spread. Changing climate parameters, slow adaptation and gene flow through seeds and pollen have to be considered as possible explanations for the late spread of the tree. Population dynamics and propagule pressure are likely to be important factors that shaped the spread of Picea abies.

The aerial plant body is derived from undifferentiated cells in the shoot apical meristem that in Arabidopsis thaliana is active throughout the plant life cycle. Upon transition to flowering the activity of the meristem is altered and the meristem starts to produce secondary inflorescences and floral meristems instead of leaves. Both the activity of the meristem and the decision of when to flower are processes strictly regulated by several mechanisms. In this thesis I describe the function of two genes that are active in the regulation of meristem function and in the regulation of when to shift to reproductive development. First, the Arabidopsis gene encoding TERMINAL FLOWER2 (TFL2), homologous to HETEROCHROMATIN PROTEIN1, was isolated and characterised. Mutations in TFL2 result in plants that are dwarfed, flowers early, have reduced sensitivity to day length and terminate the inflorescence in an apical flower. As homologues from other organisms TFL2 is active in gene regulation by gene repression. I show that the gene affect flowering time by the autonomous and the photoperiod pathways, two of four floral inductive pathways. TFL2 act to repress the activity of genes that are promoters of floral meristem identity and interacts genetically with factors known to alter the chromatin state. Further tfl2 is shown to have altered levels of and response to auxin. All together this shows that TFL2 is active as a regulator of several different processes during plant development. Second, I have characterised and studied the function of three genes encoding APETALA2 LIKE proteins in Norway spruce (Picea abies). In spruce these genes are expressed in meristems and reproductive tissues. When constitutively expressed in Arabidopsis two of the genes delays flowering time and alter the function of shoot apical and floral meristems. Together this suggests a function similar to the Arabidopsis homologues.

Norway spruce (Picea abies L. Karst) is a conifer belonging to the group gymnosperms and is an ecologically and economically important species in several parts of Europe. It is crucial for trees like Norway spruce to adapt timing of events such as bud set and growth cessation to the local environment in order to maximize the growth period while avoiding frost damage. This thesis aims at widening the knowledge about genetic control of annual growth rhythm in Norway spruce and particularly the control of bud set. Using spruce transformants ectopically expressing PaFT/TFL1-LIKE 2 (PaFTL2) the prior hypothesis that PaFTL2 induces bud set is confirmed. This is further supported by spatial and temporal expression patterns in seedlings and adult trees. It is further shown that gymnosperms possess at least two FLOWERING LOCUS T/TERMINAL FLOWER 1 (FT/TFL1)-like genes with TFL1-like function, suggesting the ancestor of FT and TFL1 to be more TFL1-like. PaFTL1 appears to have complementary expression patterns to that of PaFTL2 both spatially and temporally indicating they may act together to control growth in Norway spruce. Since bud set is controlled by photoperiod and circadian clock genes are implicated in this process, putative clock homologs were studied to gain insight into the circadian clock in gymnosperms. Several clock homologs were identified and their expression showed a diurnal pattern but the expression was rapidly damped in constant conditions. Transgenic Arabidopsis expressing putative core clock genes from spruce indicate that at least three genes, PaCCA1, PaGI and PaZTL, appear to have a conserved function between angiosperms and gymnosperms. Taken together these results suggest that gymnosperms have a similar core clock structure as angiosperms even though fundamental differences might exist since the cycling of the clock genes were rapidly damped in free-running conditions. The studies presented in this thesis support substantial conservation of pathway components controlling photoperiodic responses in angiosperms and gymnosperms and identify PaFTL2 as a component of growth rhythm control. However, important changes in these processes are also evident. The results provide a solid basis for future research on molecular mechanisms controlling an adaptive trait in an important non-model organism.

Abies is the second largest genus of family Pinaceae (after Pinus), consisting of about 51 species, all native to the Northern Hemisphere. There are six native taxa belonging to this genus growing in pure and mixed stands in Turkey. Abies cilicica subsp. isaurica, Abies nordmanniana subsp. bornm&uuml / lleriana, Abies nordmanniana subsp. equi-trojani, Abies x olcayana are endemic and considered as lower risk (LR) species. To determine the genetic relationships in Turkish firs, 18 populations of different subspecies of Abies were collected from different regions of Turkey and non-coding trn regions of chloroplast DNA were sequenced to assess the genetic structure of the studied species. trnL, trnF and trnV region were examined. All the trn regions of Abies species in the world (aproximately 300 species that found in the IPNI (The International Plant Names Index) were investigated in the database of NCBI. The available trn sequences of 23 Abies species worldwide included into the analyses. All analyses to estimate molecular diversity parameters were carried out with the MEGA software. The constructed phylogenetic tree with the trn sequences revealed that Turkish firs formed a monophyletic group with almost no sequence divergence. v Since sequence data for all three sectors of trn were not available from the NCBI data base, the phylogentic analysis with the sequence data of trnL regions were compartively analyzed in all firs. The results showed that Turkish- European species formed a single clade, which clearly differentiated them from the others, such as Japanese species, A. veitchii. Similarly, according to the sequence data of trnF, Turkish fir species were grouped together and distinctly separated from Asian-American Fir species. The results suggest that all Turkish firs may have evolved from single ancestral fir species, most likely from Abies nordmanniana.

Pineacea is the largest family of conifers that includes 51 species of Abies which is the second largest genus after Pinus. There are six native taxa in Turkey belonging to this genus. Four of these taxa (Abies cilicica subsp. isaurica, Abies nordmanniana subsp. bornm&uuml / lleriana, Abies nordmanniana subsp. equi-trojani, Abies x olcayana) are endemic and considered as low risk (LR) species according to the IUCN criteria. To determine the phylogenetic relationship in Abies spp. in Turkey, 18 populations of different taxa were collected from their natural distribution areas in Turkey. The matK gene regions of chloroplast DNA (cpDNA) were studied comparatively to reveal the genetic relationship among Turkish fir species. The available sequences from the NCBI database for the matK region of the other Abies species in the world were also investigated v comparatively with the sequences from Turkish firs. With the matK sequence data, a phylogenetic tree was constructed for the fir species and the molecular diversity parameters such as conserved and variable sites, nucleotide diversity, and evolutionary divergence were estimated using the MEGA software. The results indicated that there are no variable sites among Turkish firs with regard to matK regions of cpDNA. It appears that the matK region of cpDNA for Turkish firs is highly conserved. Since sequence data for all matK regions were not available from the NCBI data base, the phylogentic analysis with the sequence data of matK were comparatively analyzed in all firs including Turkish firs. According to matK1 region, the results showed that there were three major clades. One of the clades included all Turkish fir taxa and one species from European firs, A.numidica / however, A.holopylla, A.firma, A.veitchii, A.sachalinensis, A.nephrolepis, A.lasiocarpa, A.koreana, A.homolephis, A.fraseri, A.fargesii, A.sibirica and A.fabri formed in another clade. In addition to this, A.mariesii, A.hidalgensis, A.bracteata, A.alba formed in different major clade. According to matK1 region the results showed that Turkish firs and European firs are closer to each other. Furthermore, based on matK2 region, the results indicated that Turkish firs formed a monophyletic group.The other fir species, with respect to matK2 regions, formed different clades from Turkish firs. The results based on matK region suggest that all Turkish firs may have evolved from single ancestral fir species and the matK gene region appears to be highly conserved region in fir species.

Thesis (M.S.)--Oregon State University, 2003. / Typescript (photocopy). Includes bibliographical references. Also available on the World Wide Web.

Diss. (sammanfattning) Alnarp : Sveriges lantbruksuniversitet, 2009. / Härtill 5 uppsatser.