Onset of flowering is a major transition in the plant life cycle and is controlled by environmental factors including photoperiod, light quality and temperature. Prevalence of controlling factors depends on species, hence physiological models were selected for their strong requirement for one or another environmental cue. Among Brassicaceae, Sinapis alba was intensively studied for its strong response to photoperiod while molecular-genetic analyses of Arabidopsis thaliana disclosed complex interactions between pathways inducing flowering in response to photoperiod and other environmental cues, such as vernalization. We were therefore interested in studying the vernalization process in S. alba and its interactions with the previously characterized floral response to long days (LDs). Two-week old seedlings grown in non-inductive short days (SDs) were vernalized at 7°C for increasing durations and a strong promotive effect of vernalization was observed. In contrast to the common view of vernalization as a preliminary step bringing the competence to flower, we observed that vernalization had a direct inductive effect on flowering: floral buds were initiated during cold-exposure. Floral integrator genes SaMADSA (homologous to SUPPRESSOR OF OVEREXPRESSION OF CO 1) and SaLEAFY were up-regulated in the shoot apex after 3-4 weeks of vernalization. To monitor the vernalization process at the molecular level, we isolated SaFLC which, based on sequence analysis, expression patterns and complementation test, appeared as orthologous to FLOWERING LOCUS C. Down-regulation of SaFLC by vernalization was fast since transcript level was already very low after one week of vernalization, but stability of the repression required longer exposure to cold. To test the physiological significance of these observations, we studied the floral response to 16-h LDs after unstable and stable repression of SaFLC. We observed that one week of vernalization which was sufficient for SaFLC repression but not for maintenance of the silenced state - increased the flowering response of S. alba to LDs when the LDs just followed the cold treatment. This effect was lost after two weeks post-vernalization. In contrast, the promotive effect of longer vernalization on flowering response to LDs was maintained post-vernalization. These results suggested that vernalization not only works when plants experience long exposure to cold in winter: short cold periods might stimulate flowering of LD-plants if occurring when photoperiod is increasing, i.e. in spring.

This is a genetic study of flowering time in cultivated barley with the aim to identify the alleles contributing to rapid flowering and frost resistance. We have genotyped a collection of 23 historic barley varieties for the crucial genes [VRN-1, VRN-2, VRN-3 (HvFT), Ppd-H1, CO, and Vrs1]. We have amplified the polymorphic mutations by PCR-based methods, and sequenced them to identify possible haplotype groups. The row type was not determined of all accessions, but all the Scandinavian varieties were found to carry mutant alleles of Vrs1, that indicates them to be six-row barleys. The deletion of the crucial segment of VRN-1 vernalization contributes dominant spring growth habit. We found haplotype groups 2 and 4 to be dominant in Northern barleys whereas haplotype groups 1 and 5 dominated in south. The presence of dominant allele VRN-2 gene is addressed to floral repression until plants get vernalized. Most of the 23 varieties were found to have deleted allele of VRN-2, which is connected with a spring growth habit. The only four of the accessions that have the dominant allele of Ppd-H1 that contribute flowering are generally from the south of Europe. HvFT and CO genes CO-interact to influence flowering time. CO haplotype grouping suggest a geographical distribution of different alleles but needs more disseminations. Certain HvFT alleles cause extremely early flowering during apex development in the varieties that have deletion of VRN-2 alleles under long days. VRN-3 alleles of 14 varieties were identified.

Made available in DSpace on 2016-08-12T19:18:32Z (GMT). No. of bitstreams: 1 RafaellaRML_TESE.pdf: 8295790 bytes, checksum: d869daa8e386857296c93d9776704d7e (MD5) Previous issue date: 2015-02-24 / Conselho Nacional de Desenvolvimento Científico e Tecnológico / Garlic is one of the vegetables of great economic and social relevance in Brazil. Among the consumer states, Rio Grande do Norte, especially, despite having regions with favorable conditions for growing garlic, currently, depends on imports of this product to meet its domestic demand. The introduction of more productive and quality cultivars, better market acceptance, and adjustments of vernalization technology, a key issue for adaptation of new cultivars, are mechanisms that can contribute to garlic revitalization in areas previously producing this vegetable. In order to evaluate the development, production and quality semi-noble garlic cultivars under different periods of pre-planting vernalization of seed bulbs in two city in the Mesoregion West Potiguar, we developed two experiments, simultaneously in Baraúna and Governador Dix-sept Rosado, between the months of April and November 2012. The experimental design was a randomized complete block design with four replications. The treatments were arranged in a split plot, being the plots represented by semi-noble cultivars Gigante do Núcleo and BRS Hozan and subplots consisted by periods of vernalization before planting the seed bulbs at 4 ± 1 ° C: 0, 10, 20 and 30 days. We evaluated: plants emergency, plant height, leaf number, cultural cycle, final stand, mass fresh total, percentage of bulbs differentiation of bulbils, bulbils number, classification of bulbils, bulb average weight, yield total, classification bulbs, diameter of bulb, pH, titratable acidity, soluble solids, total soluble sugars, reducing sugars, pungency, total solids and industrial index. The vernalization did not provide increases significant in productivity, however, the use of 10 day of the vernalization pre-planting seed bulbs allowed in improving the quality of garlic semi-noble produced in Baraúna and Governador Dix-sept Rosado. The cultivars Gigante do Núcleo (4.56 t ha-1) and BRS Hozan (4.42 t ha-1) had higher yields in Baraúna. In this city, garlic produced showed greater diameter, pungency and total solids, with higher industrial index therefore better quality for industrialization (processing or dehydration) / O alho é uma das hortaliças de grande relevância econômica e social no Brasil. Entre os estados consumidores, o Rio Grande do Norte, especialmente, apesar de apresentar regiões com condições favoráveis ao cultivo de alho, atualmente, depende da importação deste produto para atender a sua demanda interna. A introdução de cultivares mais produtivas e de qualidade, com melhor aceitação de mercado, e ajustes da tecnologia de vernalização, ponto fundamental para adaptação de novas cultivares, são mecanismos que podem contribuir para revitalização do alho em regiões, anteriormente produtoras desta hortaliça. Com o objetivo de avaliar o desenvolvimento, produção e qualidade de cultivares de alho semi-nobre, submetidas a diferentes períodos de vernalização pré-plantio dos bulbos-semente em dois municípios da Mesorregião Oeste Potiguar, desenvolveram-se dois experimentos, simultaneamente, em Baraúna e Governador Dix-sept Rosado, entre os meses de abril e novembro de 2012. O delineamento experimental foi o de blocos completos casualizados com quatro repetições. Os tratamentos foram dispostos em parcelas subdivididas, sendo as parcelas representadas pelas cultivares semi-nobres Gigante do Núcleo e BRS Hozan e as subparcelas constituídas pelos períodos de vernalização pré-plantio dos bulbos-semente à 4 ± 1 ºC: 0, 10, 20 e 30 dias. Foram avaliados: emergência de plantas, altura de plantas, número de folhas, ciclo cultural, estande final, massa fresca total, percentagem de bulbos com diferenciação de bulbilhos, número de bulbilhos, classificação de bulbilhos, massa média de bulbos, produtividade total, classificação de bulbos, diâmetro de bulbos, pH, acidez titulável, sólidos solúveis, açúcares solúveis totais, açúcares redutores, pungência, sólidos totais e índice industrial. A vernalização não proporcionou aumentos significativos na produtividade, entretanto o uso de 10 dias de vernalização pré-plantio dos bulbos-semente possibilitou incrementos na qualidade do alho semi-nobre produzido em Baraúna e Governador Dix-sept Rosado. As cultivares Gigante do Núcleo (4,56 t ha-1) e BRS Hozan (4,42 t ha-1) apresentaram maior produtividade em Baraúna. Neste município, o alho produzido apresentou maior diâmetro, além de pungência e sólidos totais, com índice industrial mais elevado e, portanto, melhor qualidade para industrialização (processamento ou desidratação)

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / Wheat is the main winter crop in Rio Grande do Sul State. Temperature and photoperiod are the major environmental factors that control development in wheat. Wheat plants exposed to cold, non freezing temperatures induce them to enter the reproductive phase. Temperature and photoperiod response functions are used to simulate the wheat development. Developmental models are important part of water soil balance, growth and yield crop models. Thus, the objectives of this thesis were: (i) to evaluate vernalization sensitivity of some Brazilian wheat genotypes and to fit a vernalization response function for these genotypes; (ii) to determine the best temperature, photoperiod and vernalization functions to simulate wheat development with the Wang and Engel (WE) model for some Brazilian wheat genotypes; and (iii) to improve the prediction of available soil water in the soil by modifying two mechanistic models. To achieve the first objective, a factorial experiment with eight levels of vernalization (0, 7, 14, 21, 28, 35, 42 and 49 vernalization days) and six wheat genotypes were used. Also a field experiment was carried out in Santa Maria, RS, Brazil, to provide independent data on the date of developmental stages of six Brazilian wheat genotypes for evaluating the original and the modified WE model. Soil water content was determined with the gravimetric method to evaluate water balance models during three sowing dates. For those genotypes that presented some response to vernalization, response functions are presented, which can be used in models for simulating the development of Brazilian wheat cultivars with distinct vernalization requirements. The WE model simulates better the development of the Brazilian wheat genotypes using cardinal temperatures higher than original WE model and using daily maximum and minimum temperatures in the calculation of the temperature response function. The modified Campbell; Diaz water balance model is more realistic and has a greater potential for performing well in regions others than it was developed. The modified Amir; Sinclair water balance model simulates better the available soil water than the original model. / O trigo é a principal cultura de inverno do Estado do Rio Grande do Sul. A temperatura e o fotoperíodo são os dois elementos meteorológicos que mais influenciam o desenvolvimento de plantas de trigo. Quando se expõem plantas de trigo a temperaturas baixas não congelantes, estas podem ser induzidas a entrarem no estágio reprodutivo, diminuindo o seu ciclo de desenvolvimento. Para simular o desenvolvimento do trigo são utilizadas funções de resposta a temperatura e fotoperíodo. O modelo de desenvolvimento é parte importante em modelos de simulação de balanço de água no solo, crescimento e rendimento das culturas. Assim, os objetivos desta tese foram: (i) determinar a sensibilidade à vernalização de alguns genótipos de trigo do sul do Brasil e, ajustar uma função de reposta a vernalização para aqueles que apresentaram sensibilidade; (ii) determinar a melhor função de temperatura, fotoperíodo e vernalização para simular o desenvolvimento de alguns genótipos brasileiros de trigo com o modelo de Wang e Engel (WE); e (iii) obter uma melhor predição do balanço de água no solo, modificando dois modelos mecanísticos de balanço de água no solo. Para atingir o primeiro objetivo, foi conduzido um experimento fatorial com oito níveis de vernalização (0, 7, 14, 21, 28, 35, 42 e 49 dias de vernalização) e seis genótipos de trigo. Também foi realizado um experimento com várias datas de semeadura em Santa Maria, RS, Brasil, para obtenção de dados independentes dos estágios de desenvolvimento de seis genótipos Brasileiros de trigo para avaliar o modelo WE modificado. Foi determinado o conteúdo de água no solo pelo método gravimétrico em três datas de semeadura, com a finalidade de avaliar o desempenho dos dois modelos de balanço de água no solo. Para os genótipos que apresentaram resposta à vernalização são apresentadas funções de resposta, que podem ser usadas em modelos de simulação do desenvolvimento de genótipos brasileiros de trigo. O modelo WE simula melhor o desenvolvimento do trigo quando são usadas as temperaturas cardinais maiores do que as do modelo original e usando as temperaturas mínimas e máximas diária para o cálculo da função de resposta a temperatura. O modelo de balanço de água no solo de Campbell; Diaz modificado é mais realístico e com maior possibilidade de desempenho satisfatório em regiões de clima distinto daquelas em que foi desenvolvido. O modelo de balanço de água no solo de Amir; Sinclair modificado estima melhor a água disponível no solo que o modelo original.

Abstract Adaptation to local environment is important for all organisms to guarantee survival and to maximize reproduction. Populations of the same species may live in environments that differ markedly. Due to differential selection pressures this can lead to population differentiation, which can be studied both at the phenotypic and at the gene level. The growing season cued by long days is typically short in the north, whereas southern populations have long growing seasons and are adapted to short days. Seasonal fluctuations in temperature also differ between northern and southern environments. Daylength and temperature regulate the timing of flowering in plants. Environmental regulation of flowering and its genetic basis has been extensively studied in the annual model species Arabidopsis thaliana. The perennial growth and flowering habit has been studied especially in trees, but studies on herbaceous plants species have been lacking. In this thesis, I have studied adaptation to growing season length in a perennial herbaceous model species Arabidopsis lyrata. Individuals from populations adapted to northern and southern environments in Europe were grown in same conditions in the growth chambers and in the field. Differentiation between the populations was studied by observing their flowering phenotypes and by studying the expression of genes that are candidates for governing the phenotypic differentiation. The main result in the thesis was that adaptation to short growing season in north can be seen as long daylength requirement for flowering and as fast developmental rate. Critical daylength for flowering likely regulates especially the timing of flowering cessation in the end of the growing season. Flowering time of individuals from northern populations also responded more strongly to cold treatment (representing winter) than that of the southern population. The cold requirement for flowering guarantees that the plants only flower after the winter in the spring with suitable conditions. Expression studies indicated that population differentiation in flowering could be at least partly governed by the expression variation in a few candidate genes. The results in this thesis are valuable for instance for understanding perennial species in general, including tree and crop species, and for predicting how plants response to changing climate. / Tiivistelmä Ympäröiviin oloihin sopeutuminen on tärkeää kaikille organismeille selviytymisen ja jälkeläistuoton kannalta. Saman lajin eri populaatiot saattavat elää ympäristöissä, joiden olosuhteet poikkeavat toisistaan huomattavasti. Tällöin populaatioihin kohdistuvat erilaiset valintapaineet ja populaatiot erilaistuvat. Erilaistuminen havaitaan tarkastelemalla yksilöiden ilmiasuja ja geenejä. Kasvukauden pituus määrittää eteläisten ja pohjoisten kasvuympäristöjen valintapaineita. Pohjoisessa kasvukausi on lyhyt ja sen alkamisesta ja loppumisesta kertoo pitkä päivänpituus. Etelässä on pitkä kasvukausi ja siellä elävät populaatiot ovat sopeutuneet lyhyeen päivänpituuteen. Myös vuodenaikaiset lämpötilavaihtelut eroavat pohjoisten ja eteläisten alueiden välillä. Muutokset päivänpituudessa ja lämpötilassa säätelevät kasveilla kukkimisen ajankohtaa. Kukkimiseen vaikuttavia ympäristötekijöitä ja kukkimista sääteleviä geenejä on tutkittu paljon yksivuotisella mallilajilla lituruoholla (Arabidopsis thaliana). Monivuotisten kasvien kasvun ja kukkimisen säätelyä on tutkittu etenkin puilla, mutta hyvin vähän ruohovartisilla kasveilla. Tässä väitöskirjatyössä tutkin kasvukauden pituuteen sopeutumista monivuotisella ruohovartisella lajilla, idänpitkäpalolla (Arabidopsis lyrata). Pohjoisiin ja eteläisiin ympäristöoloihin sopeutuneiden eurooppalaisten populaatioiden yksilöitä kasvatettiin samanlaisissa olosuhteissa kontrolloiduissa kasvatushuoneissa ja kenttäolosuhteissa. Populaatioiden erilaistumista tarkkailtiin kukkimiseen liittyvissä ominaisuuksissa sekä eroja selittävien kandidaattigeenien ekspressiossa. Tutkimuksen päätulos oli, että pohjoisen populaation lyhyeen kasvukauteen sopeutuminen voidaan havaita pitkän päivän vaatimuksena kukkimiselle ja nopeana kehityksenä. Kenttäolosuhteissa päivänpituus sääteli etenkin kukkimisen lopetusta kasvukauden lopussa. Pitkä kylmäkäsittely nopeutti kukkimista etenkin pohjoisessa populaatiossa. Kukkimista edeltävä kylmävaatimus takaa, etteivät kasvit kuki syksyllä epäsuotuisissa olosuhteissa, vaan vasta keväällä talven jälkeen. Populaatioiden erilaistuminen kukkimisen päivänpituus- ja kylmävaatimuksessa selittyy todennäköisesti ainakin osittain kandidaattigeenien ekspressioeroilla. Tämän väitöskirjatyön tuloksia voidaan soveltaa monivuotisten viljelykasvien jalostuksessa tai ennustettaessa kasvipopulaatioiden sopeutumista ilmastonmuutokseen.

Abstract Arabidopsis lyrata is a close outcrossing relative of Arabidopsis thaliana, the model organism of plant physiology and molecular biology. I studied variation in flowering time and the factors shaping the variation within and between A. lyrata populations in different environments. The role of the two important proximate factors determining flowering time, day length and temperature, were studied in climate chambers. The southern A. lyrata populations were found to flower in high frequency and quicker than northern A. lyrata populations in all studied environments, but the reaction of northern populations on long day length was found to be stronger than that of southern populations. Differences in vernalization requirement between A. lyrata populations were found in outdoor common garden, but in the climate chambers the results of vernalization experiments were not consistent. Strength and direction of selection on flowering time and other life history traits were studied in alpine and lowland A. lyrata populations in Scandinavia. Differences in selection were found both between populations and between years. Grazing sheep caused high levels of damage in inflorescences in the alpine population. In the lowland population there was less herbivory, caused by insects and hares. The difference in selection on flowering traits in the two study populations might be partly caused by selective grazing. Completely outcrossing mating system in A. lyrata is due to well developed self-incompatibility system. However, biparental inbreeding is likely to exist in natural populations and it may lead to spatial structuring of genetic variation within populations. I studied the effects of biparental inbreeding on components of fitness in A. lyrata in three different environments. I found inbreeding depression after sib-mating to be substantial. Stressful environment reduced the overall performance of the plants, but had no effect on the magnitude of inbreeding depression. A literature survey indicates that the observed levels of inbreeding depression in self-incompatible A. lyrata were higher than those of self-compatible species. This suggests that self-compatible species have purged some of their genetic load. The genetic basis of flowering time variation in A. lyrata can be further studied by using A. thaliana molecular tools.

Jointed goatgrass (Aegilops cylindrica Host) infestations in winter wheat (Triticum aestivum L.) production regions of the central and western USA result in severe economic losses in the wheat market. Field and greenhouse studies were conducted to determine the vernalization requirements of winter wheat, spring wheat, jointed goatgrass, and wheat by jointed goatgrass reciprocal hybrid plants. In field studies, jointed goatgrass plants required more vernalization to produce spikelets and germinable seed than 'Madsen' winter wheat plants. In greenhouse studies, plants of jointed goatgrass populations collected from Oregon and Washington wheat fields required fewer vernalization days to reach the joint stage than Madsen plants. Detailed observations in the greenhouse revealed a longer period between jointing and anthesis for most jointed goatgrass populations that was overlooked in field studies. Vernalization for 6-wk represents the minimum treatment for synchronous reproductive development among jointed goatgrass populations, Madsen winter wheat, and Madsen by jointed goatgrass hybrids, yet the risk of gene transfer might be greater after 7-wk vernalization. In the jointed goatgrass populations tested, there was not selection for a vernalization insensitive growth habit. Because jointed goatgrass spikelets often contain two seed, germination was recorded for primary and secondary positioned seed. Germination of freshly harvested jointed goatgrass seed was promoted by 25/15 C day/night temperatures. However, light and 30/20 C incubation was necessary for maximum germination of non-dormant, primary positioned seed. Both primary and secondary positioned seed within jointed goatgrass spikelets were non-dormant after 16-wk after-ripening at 22 ± 2 C. Under optimum growing conditions, no planting depth selectively allowed wheat germination and emergence while preventing jointed goatgrass germination and emergence. Glume removal did not alleviate dormancy completely in jointed goatgrass seed. Research confirmed jointed goatgrass population polymorphism for vernalization requirements and seed dormancy. Jointed goatgrass reproductive variability might be part of a general purpose genotype strategy to germinate and colonize a wide range of environments. Wheat by jointed goatgrass hybrid plants should be removed from winter and spring wheat fields. Despite a short dormancy period, three or more years of rotation outside of winter annual crops may be necessary to reduce populations of jointed goatgrass. / Graduation date: 2006