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Modern crop production is characterized by high nitrogen (N) application rates, which can influence the co-limitation of harvested yield by other nutrients. Using a multidimensional niche volume concept and scaling exponents frequently applied in plant ecological research, we report that increased N and phosphorus (P) uptake in a growing wheat crop along with enhanced grain biomass is associated with more than proportional increase of other nutrients. Furthermore, N conversion efficiency and grain yield are strongly affected by the magnesium (Mg) to P ratio in the growing crop. We analyzed a field trial in Central Sweden including nine wheat varieties grown during two years with contrasting weather, and found evidence for Mg co-limitation at lower grain yields and P co-limitation at higher yields. We argue that critical concentrations of single nutrients, which are often applied in agronomy, should be replaced by nutrient ratios. In addition, links between plant P and Mg contents and root traits were found; high root number enhanced the P:N ratio, whilst steep root angle, indicating deep roots, increased the Mg:N ratio. The results have significant implications on the management and breeding targets of agriculturally grown wheat, which is one of the most important food crops worldwide.

Perennial cereal crops could limit the negative impacts of agriculture on the environment and climate change. In cold temperate climates, perennial plants must be adapted to seasonal changes and abiotic stresses, such as frost, to be able to regrow for several years. Wild crop relatives that are perennials and already adapted to cold temperate climates may provide genetic resources for breeding new perennial cereal grain crops. Barley (Hordeum vulgare) is one of the most important cereals in northern agricultural areas, and its related perennial species may be good candidates for the development of perennial cereals. We evaluated a diverse set of 17 wild perennial Hordeum species represented by 67 accessions in field conditions with a cold winter climate and long days during summer in Central Sweden (latitude 60&deg, N). Six species (H. brevisubulatum, H. bulbosum, H. fuegianum, H. jubatum, H. lechleri and H. secalinum) showed regrowth and formation of spikes for four seasons. The most distant perennial relative of barley, H. stenostachys, showed weak regrowth. H. bulbosum, the closest perennial barley relative, had a large number of accessions with wide geographic origins that showed good regrowth. Together with its storage bulbs and its cross-compatibility with barley, this makes H. bulbosum an important genetic resource for the development of perennial Hordeum grains using either the domestication or the wide-hybridization strategy.

A network of 21 experiments was established across nine countries on four continents and spanning both hemispheres, to evaluate the relative performance of early generation perennial cereal material derived from wheat, rye, and barley and to inform future breeding strategies. The experimental lines were grown in replicated single rows, and first year production and phenology characteristics as well as yield and persistence for up to three years were monitored. The study showed that the existing experimental material is all relatively short-lived (≤3 years), with environments that are milder in summer and winter generally conferring greater longevity. No pedigree was superior across this diverse network of sites although better performing lines at the higher latitude sites were generally derived from Thinopyrum intermedium. By contrast, at lower latitudes the superior lines were generally derived from Th. ponticum and Th. elongatum parentage. The study observed a poor relationship between year 1 performance and productivity in later years, highlighting the need for perennial cereal material with greater longevity to underpin future experimental evaluation, and the importance for breeding programs to emphasize post-year 1 performance in their selections. Hybrid lines derived from the tetraploid durum wheat generally showed greater longevity than derivatives of hexaploid wheat, highlighting potential for greater use of Triticum turgidum in perennial wheat breeding. We advocate a model in future breeding initiatives that develops perennial cereal genotypes for specific target environments rather than a generic product for one global market. These products may include a diversity of cultivars derived from locally adapted annual and perennial parents. In this scenario the breeding program may have access to only a limited range of adapted perennial grass parents. In other situations, such as at very high latitude environments, perennial crops derived from barley or rye may have a better chance of success than those derived from wheat. In either case, development and selection of the perennial parent for adaptation to local environments would seem fundamental to success.

Albinism is a major limitation in the production of doubled haploid plants in cereals. As this trait is partly genetically controlled, the culture method has to be adapted to the plant genotype to reduce albinism. We have improved green plant recovery in the Northern barley ( Hordeum vulgare L.) spring cultivar Mitja from 6 to 42% by altering the culture method and composition of the culture media. In a new three-step protocol, the addition of casein hydrolysate to an inter - mediate regeneration medium gave the high - est number of green plants. The shoot and root growth medium used for germinated embryos was superior to the standard media reported in the literature for root formation and shoot growth. o ur modified method has merit for test - ing with other recalcitrant genotypes in anther and isolated microspore cultures.

The efficient use of nutrients underpins food security and reduces losses of nutrients to the environment. Different approaches have been applied to improve and evaluate the nutrient use efficiency (NUE) of plants, depending on the purpose to which the concept has been put. The various approaches differ greatly in the scale and target for NUE improvement and assessment. In this chapter, various approaches and conceptions for improving and evaluating NUE across different scales are discussed: (1) the investigation of the physiology and genetics of NUE at single tissue, cell and molecular scales, targeting understanding and improvement of NUE-related traits through, for example, plant breeding; and (2) the assessment of NUE and nutrient balances at whole plant, field, and regional scales for evaluating crops with NUE-improved traits. Appropriate methodologies are available to link the processes acting at molecular, individual plant, farm, and regional scales, but require increased collaborative efforts between the different disciplines.

Cassava ranks fifth among crops in global starch production. It is used as staple food in many tropical countries of Africa, Asia and Latin America. In (the) People’s Republic of China, although not yet a staple food, cassava is of major economic importance for starch for a large area of southern (the) PRC, especially in the provinces of Guangdong, Guanxi, Yunnan and Hainan. Recently, cassava-derived bioethanol production has been increasing due to its economic benefits compared to other bioethanol-producing crops in the country. We discuss here the possible potentials of cassava for bioethanol production.

Cassava (Manihot esculenta Crantz) is a starchy root crop grown in the tropics mainly by small-scale farmers even though agro-industrial processing is rapidly increasing. For this processing market improved varieties with high dry matter root content (DMC) is required. Potentially toxic cyanogenic glucosides are synthesized in the leaves and translocated to the roots. Selection for varieties with low cyanogenic glucoside potential (CNP) and high DMC is among the principal objectives in cassava breeding programs. However, these traits are highly influenced by the environmental conditions and the genetic control of these traits is not well understood. An S1 population derived from a cross between two bred cassava varieties (MCOL 1684 and Rayong 1) that differ in CNP and DMC was used to study the heritability and genetic basis of these traits. A broad-sense heritability of 0.43 and 0.42 was found for CNP and DMC, respectively. The moderate heritabilities for DMC and CNP indicate that the phenotypic variation of these traits is explained by a genetic component. We found two quantitative trait loci (QTL) on two different linkage groups controlling CNP and six QTL on four different linkage groups controlling DMC. One QTL for CNP and one QTL for DMC mapped near each other, suggesting pleiotrophy and/or linkage of QTL. The two QTL for CNP showed additive effects while the six QTL for DMC showed additive effect, dominance or overdominance. This study is a first step towards developing molecular marker tools for efficient breeding of CNP and DMC in cassava.

Cassava is a tropical crop and grown for its tuberous starchy roots. In Africa it is mainly cultivated by small-scale farmers who observe, select and name their cassava varieties based on morphology, food, social and economic interest. Here we have used an interdisciplinary approach involving farmer interviews, genetic markers and morphological descriptors to study the composition of cassava varieties on small-scale farms in 11 villages located in three districts in Uganda, the genetic structure within and between these varieties and their morphology. The composition of local, newly introduced and improved varieties differed widely between villages and districts. The Ugandan farmers in our study seemed to adopt improved varieties to a greater extent when there was a nearby market, prevalence of disease epidemics and good extension service. We found considerable genetic variation both within and between cassava varieties though the variation was larger between varieties. However, most local and improved varieties showed predominating genotypes at many loci. Accessions of commonly grown varieties meeting farmers' preferences could therefore be selected and implemented in future breeding programmes involving development, dissemination and adoption. The like-named varieties in different villages were genetically similar, demonstrating farmers' ability to differentiate and maintain the same variety over large areas. However, some varieties with different names in different villages showed both genetic and morphological similarity, suggesting that farmers may rename plants when they are introduced into their fields. The large differences found in variety and genetic composition between villages and districts in Uganda may be a result of the diverse needs and growing conditions characteristic for traditional farming system. This suggests that efforts to conserve and increase the genetic diversity in farmers' fields will require policies tailored to each area.

We used quantitative trait locus/loci (QTL) mapping to study the inheritance of traits associated with perennialism in a cross between an annual (Zea mays ssp. parviglumis) and a perennial (Z. diploperennis) species of teosinte. The most striking difference between these species is that Z. diploperennis forms rhizomes, whereas Z. mays ssp. parviglumis lacks these over-wintering underground stems. An F2 population of 425 individuals was genotyped at 95 restriction fragment length polymorphism marker loci and the association between phenotype and genotype was analyzed by composite interval mapping. We detected a total of 38 QTL for eight traits. The number of QTL found for each trait ranged from two for rhizome formation to nine for tillering. QTL for six of the traits mapped near each other on chromosome 2, and QTL for four traits mapped near each other on chromosome 6, suggesting that these regions play an important role in the evolution of the perennial habit in teosinte. Most of the 38 QTL had small effects, and no single QTL showed a strikingly large effect. The map positions that we determined for rhizome formation and other traits in teosinte may help to locate corresponding QTL in pasture and turf grasses used as forage for cattle and for erosion control in agro-ecosystems.

Seed predator/pollinator and host plant interactions, which may be considered as antagonistic, have the potential to provide good model systems for the study of the early stages of evolution towards mutualism. I describe a relationship between a seed predator, the geometrid moth Perizoma affinitatum, and the dioecious plant Silene dioica. The moth is an obligate seed predator on its host plant. The searching and ovipositing behaviour of the female moths, number of eggs deposited per flower, the pollinating ability of the moths and the seed consumption by the larvae are described as different parameters and studied in two Finnish coastal populations. A high pollinating ability and limited seed consumption by the predator was found and discussed in relation to fitness models of P. affinitatum and S. dioica. In a mutualistic relationship there must be a balance between the costs and benefits so that the seed production by the moths is larger than the seed consumption by the larvae, given a net seed output larger than zero. The data of the parameters included in a seed production/consumption model give a positive seed output when the proportion of S. dioica flowers pollinated by other non-predating insects is less than 60%. Accordingly, even if P. affinitatum would become the exclusive pollinator it would not endanger the survival of the host plant and both partners would benefit from this interaction. Limited seed consumption, high pollinating ability and host specificity as seen in the P. affinitatum and S. dioica interaction are considered to have been important pre-existing qualities in the evolution of the obligate mutualisms between yucca and yucca moths and fig and fig wasps. In isolated serpentine populations where the gene flow is restricted and co-pollinators are rare the interaction between P. affinitatum and S. dioica has the potential to shift from parasitism to mutualism.

Summary •The effects of Ni and Mg, two factors involved in the infertility of serpentine soils, were studied in the alpine plant Cerastium alpinum. Root growth of plants from adjacent serpentine and non-serpentine populations in Scandinavia, representing an eastern and western postglacial immigration lineage and the hybrid zone between them, were compared to study the adaptation of C. alpinum populations. •Seedlings were placed in solutions with low or high concentrations of Ni and Mg in a full factorial experiment according to a randomized block design. The growth of roots was analyzed and discussed in relation to the soil content. •The serpentine populations showed higher tolerance to Ni and Mg stress than non-serpentine populations. The degree of metal tolerance differed among the serpentine populations and was related to the effective concentrations of Ni and Mg in the soil at each site. •The results suggest that serpentine tolerance is locally evolved in C. alpinum and that tolerance has arisen in parallel during the postglacial colonization of Scandinavia on serpentine soils with similar composition.

We investigated the variation in plant response in host-pathogen interactions between wild (Aegilops spp., Triticum spp.) and domesticated wheat (Triticum spp.) and Wheat dwarf virus (WDV). The distribution of WDV and its wild host species overlaps in Western Asia in the Fertile Crescent, suggesting a coevolutionary relationship. Bread wheat originates from a natural hybridization between wild emmer wheat (carrying the A and B genomes) and the wild D genome donor Aegilops tauschii, followed by polyploidization and domestication. We studied whether the strong selection during these evolutionary processes, leading to genetic bottlenecks, may have resulted in a loss of resistance in domesticated wheat. In addition, we investigated whether putative fluctuations in intensity of selection imposed on the host-pathogen interactions have resulted in a variation in susceptibility to WDV. To test our hypotheses we evaluated eighteen wild and domesticated wheat taxa, directly or indirectly involved in wheat evolution, for traits associated with WDV disease such as leaf chlorosis, different growth traits and WDV content. The plants were exposed to viruliferous leafhoppers (Psammotettix alienus) in a greenhouse trial and evaluated at two time points. We found three different plant response patterns: i) continuous reduction in growth over time, ii) weak response at an early stage of plant development but a much stronger response at a later stage, and iii) remission of symptoms over time. Variation in susceptibility may be explained by differences in the intensity of natural selection, shaping the coevolutionary interaction between WDV and the wild relatives. However, genetic bottlenecks during wheat evolution have not had a strong impact on WDV resistance. Further, this study indicates that the variation in susceptibility may be associated with the genome type and that the ancestor Ae. tauschii may be useful as genetic resource for the improvement of WDV resistance in wheat.

Background Wild barley is adapted to highly diverse environments throughout its geographical distribution range. Transcriptome sequencing of differentially adapted wild barley ecotypes from contrasting environments contributes to the identification of genes and genetic variation involved in abiotic stress tolerance and adaptation. Results Two differentially adapted wild barley ecotypes from desert (B1K2) and Mediterranean (B1K30) environments were analyzed for drought stress response under controlled conditions. The desert ecotype lost more water under both irrigation and drought, but exhibited higher relative water content (RWC) and better water use efficiency (WUE) than the coastal ecotype. We sequenced normalized cDNA libraries from drought-stressed leaves of both ecotypes with the 454 platform to identify drought-related transcripts. Over half million reads per ecotype were de novo assembled into 20,439 putative unique transcripts (PUTs) for B1K2, 21,494 for B1K30 and 28,720 for the joint assembly. Over 50% of PUTs of each ecotype were not shared with the other ecotype. Furthermore, 16% (3,245) of B1K2 and 17% (3,674) of B1K30 transcripts did not show orthologous sequence hits in the other wild barley ecotype and cultivated barley, and are candidates of ecotype-specific transcripts. Over 800 unique transcripts from each ecotype homologous to over 30 different stress-related genes were identified. We extracted 1,017 high quality SNPs that differentiated the two ecotypes. The genetic distance between the desert ecotype and cultivated barley was 1.9-fold higher than between the Mediterranean ecotype and cultivated barley. Moreover, the desert ecotype harbored a larger proportion of non-synonymous SNPs than the Mediterranean ecotype suggesting different demographic histories of these ecotypes. Conclusions The results indicate a strong physiological and genomic differentiation between the desert and Mediterranean wild barley ecotypes and a closer relationship of the Mediterranean to cultivated barley. A significant number of novel transcripts specific to wild barley were identified. The higher SNP density and larger proportion of SNPs with functional effects in the desert ecotype suggest different demographic histories and effects of natural selection in Mediterranean and desert wild barley. The data are a valuable genomic resource for an improved genome annotation, transcriptome studies of drought adaptation and a source of new genetic markers for future barley improvement. Electronic supplementary material The online version of this article (doi:10.1186/1471-2164-15-995) contains supplementary material, which is available to authorized users.

The mutualisms between fig trees and their pollinator fig wasps and between yucca plants and yucca moths are spectacular examples of coevolution. The characteristics of these independently evolved mutualisms have resulted from long-term processes, the first stages of which are unknown. A fundamental question in the study of mutualism is how these interactions evolve. Seed predator/pollinator and host plant interactions, which may initially be considered as mainly antagonistic, have the potential to provide good model systems for the study of the first stages of evolution towards mutualism. We present here theoretical models assessing the consequences of interactions between specialized seed predator insects and their host plants. These models describe the parameters that affect the fitness of an individual female seed predator and her influence on the fitness of the host plant. In an optimal strategy for the seed predator, the number of eggs laid in each flower depends on the interaction between the adult and larva survival. Along with a growing predation pressure on adults and larvae several eggs must be laid in each flower by the female seed predator to enhance her fitness. However, in a situation where the host plant selectively aborts flowers with a high number of eggs the fitness of the seed predator will seriously decrease. If the cost of selective abortion is less than the cost of seed predation the host plant will maintain fitness. In a mutualistic relationship a balance between the cost and the benefit of the parameters in the fitness models of the seed predator and the host plant has to occur so that the net seed output is larger than zero (0). Any unselfish behaviour or quality of the seed predator that would benefit the host plant in such a way that the net seed output increases might be a first stage in an interaction becoming mutualistic. The models presented here will not only provide a platform for empirical studies on interactions that may swing from parasitism to mutualism, but also for seed predator/pollinator and host plant interactions in general.

Some Swedish spring wheat varieties have recently been shown to carry a rare wildtype (wt) allele of the gene NAM-B1, known to affect leaf senescence and nutrient retranslocation to the grain. The wt allele is believed to increase grain protein concentration and has attracted interest from breeders since it could contribute to higher grain quality and more nitrogen-efficient varieties. This study investigated whether Swedish varieties with the wt allele differ from varieties with one of the more common, non-functional alleles in order to examine the effect of the gene in a wide genetic background, and possibly explain why the allele has been retained in Swedish varieties. Forty varieties of spring wheat differing in NAM-B1 allele type were cultivated under controlled conditions. Senescence was monitored and grains were harvested and analyzed for mineral nutrient concentration. Varieties with the wt allele reached anthesis earlier and completed senescence faster than varieties with the non-functional allele. The wt varieties also had more ears, lighter grains and higher yields of P and K. Contrary to previous information on effects of the wt allele, our wt varieties did not have increased grain N concentration or grain N yield. In addition, temporal studies showed that straw length has decreased but grain N yield has remained unaffected over a century of Swedish spring wheat breeding. The faster development of wt varieties supports the hypothesis of NAM-B1 being preserved in Fennoscandia, with its short growing season, because of accelerated development conferred by the NAM-B1 wt allele. Although the possible effects of other gene actions were impossible to distinguish, the genetic resource of Fennoscandian spring wheats with the wt NAM-B1 allele is interesting to investigate further for breeding purposes.

Serpentine soils are hostile to plant life. They are dry, contain high concentrations of nickel and have an unfavorable calcium/magnesium ratio. The dioecious plant Silene dioica (L.) Clairv. (Caryophyllaceae) is the most common herb on serpentine soils in the Swedish mountains. It also commonly grows on non-serpentine soils in the subalpine and coastal area. I have compared the germination frequency, plant establishment and growth of serpentine and subalpine non-serpentine populations in serpentine soil under greenhouse conditions. Further more I have studied the specific effect of nickel on root and shoot growth of serpentine and non-serpentine plants from the subalpine and coastal area in solutions with different concentrations of nickel. Plants from serpentine and non-serpentine populations grew well and in a similar fashion in serpentine soil. Moreover, S. dioica plants, irrespective of original habitat, tolerated enhanced concentrations of nickel when grown in solutions. An analysis of metal content in serpentine plants from natural populations shows that S. dioica has a higher nickel concentration in the roots than in the shoots. The growth studies show that S. dioica is constitutively adapted to serpentine, and that all populations have the genetic and ecological tolerance to grow on serpentine.

Endemic Hawaiian Silene colonize new lava flows and are outcompeted as the ground ages. They illustrate the genetic processes operating in the evolution of pioneering island plants. The volcanic history of the Hawaiian Islands allows an estimation of the age of these plant populations. In this study, populations of S. struthioloides from Maui and the older part of the island of Hawaii, and S. ha aliensis from the youngest volcanoes of the island of Hawaii were analyzed by enzyme electrophoresis. The genetic structures of these populations were placed in a geographic and geologic context. Silene is much more polymorphic on the older island, Maui. Genetic variation appears to have been lost in the colonization of the youngest island, Hawaii. Interestingly, some loci for populations on the younger volcanoes of the island of Hawaii are monomorphic for alleles not found in populations on the older part of Hawaii. Recurrent colonizations are accompanied with founder effects, and restricted gene flow among populations has led to isolation and genetic drift. This has resulted in genetic and morphological differentiation of Silene populations on the youngest volcanoes.

Cassava (Manihot esculenta Crantz) is a starchy root crop grown in the tropics mainly by small-scale farmers even though agro-industrial processing is rapidly increasing. For this processing market improved varieties with high dry matter root content (DMC) is required. Potentially toxic cyanogenic glucosides are synthesized in the leaves and translocated to the roots. Selection for varieties with low cyanogenic glucoside potential (CNP) and high DMC is among the principal objectives in cassava breeding programs. However, these traits are highly influenced by the environmental conditions and the genetic control of these traits is not well understood. An S(1) population derived from a cross between two bred cassava varieties (MCOL 1684 and Rayong 1) that differ in CNP and DMC was used to study the heritability and genetic basis of these traits. A broad-sense heritability of 0.43 and 0.42 was found for CNP and DMC, respectively. The moderate heritabilities for DMC and CNP indicate that the phenotypic variation of these traits is explained by a genetic component. We found two quantitative trait loci (QTL) on two different linkage groups controlling CNP and six QTL on four different linkage groups controlling DMC. One QTL for CNP and one QTL for DMC mapped near each other, suggesting pleiotrophy and/or linkage of QTL. The two QTL for CNP showed additive effects while the six QTL for DMC showed additive effect, dominance or overdominance. This study is a first step towards developing molecular marker tools for efficient breeding of CNP and DMC in cassava.

The mode of inheritance of six enzyme markers in the octoploid alpine plant Cerastium alpinum was analyzed. Offspring from crosses between heterozygotes showed fixed heterozygosity at malate dehydrogenase-2, phosphoglucoisomerase-2, triosephosphate isomerase-2, and triosephosphate isomerase-3. Phosphoglucomutase-1 also showed fixed heterozygosity except in offspring from one cross. Fixed heterozygosity in five enzyme systems suggests that C. alpinum has originated through at least some allopolyploidization. Offspring from plants heterozygous for two alleles at the menadione reductase-1 (Mr-1) locus did not deviate significantly from a 1:2:1 ratio. The large proportion of homozygotes suggests disomic inheritance because any kind of polysomic inheritance would result in a substantially increased proportion of heterozygotes relative to disomic inheritance. Assuming a diploid model for Mr-1, this locus was used to analyze the population genetic structure within C. alpinum populations. Inbreeding was found in many alpine populations. This may help explain the large genetic distances found among alpine populations in a previous study. The analysis is only based on one segregating locus, and the results should therefore be treated with caution. However, by establishing the mode of inheritance through crosses, we have been able to use a codominant marker in population genetic analysis of an octoploid plant.

Cassava (Manihot esculenta) is a tropical crop that is grown in Africa, Latin America and Southeast Asia. Cassava was introduced from Latin America into West and East Africa at two independent events. In Uganda a serious threat to cassava's survival is the cassava mosaic disease (CMD). Uganda has had two notable CMD epidemics since the introduction of cassava in the 1850s causing severe losses. SSR markers were used to study the effect of CMD on the genetic diversity in five agroecologies in Uganda with high and low incidence of CMD. Surprisingly, high gene diversity was detected. Most of the diversity was found within populations, while the diversity was very small among agroecological zones and the high and low CMD incidence areas. The high genetic diversity suggests a mechanism by which diversity is maintained by the active involvement of the Ugandan farmer in continuously testing and adopting new genotypes that will serve their diverse needs. However, in spite of the high genetic diversity we found a loss of rare alleles in areas with high CMD incidence. To study the effect of the introgression history on the gene pool the genetic differentiation between East and West Africa was also studied. Genetic similarities were found between the varieties in Uganda and Tanzania in East Africa and Ghana in West Africa. Thus, there is no evidence for a differentiation of the cassava gene pool into a western and an eastern genetic lineage. However, a possible difference in the genetic constitution of the introduced cassava into East and West Africa may have been diminished by germplasm movement.

To investigate the genetic mechanisms that underlie morphological evolution in natural populations, we employed QTL mapping to dissect the inheritance of leaf sheath characters that distinguish Chalco from Balsas teosinte. Abundant macrohairs (trichomes) and intense anthocyanin accumulation are found in Chalco teosinte sheaths whereas Balsas teosinte leaf sheaths are green and glabrous. These character states may represent adaptations to the cooler highland (Chalco) vs. warmer middle-elevation (Balsas) climates. QTL mapping in multiple populations revealed a mix of major- and minor-effect QTL affecting both sheath color (anthocyanin) and macrohair abundance. The major QTL for macrohairs accounts for 52% of the parental difference. Epistatic interactions were detected between the major-effect QTL and multiple other QTL for both traits, accounting for substantial portions of phenotypic variance. Developmental analyses suggest that regulatory program changes underlie the phenotypic differences. Sheath anthocyanin QTL are clearly associated with b1 and a3, both of which are regulators of anthocyanin biosynthesis. Our findings suggest that changes in a small number of QTL can lead to morphological evolution by modulating existing developmental programs.

The plant cover of Fennoscandia is young because of the recent glaciation. This study covers the early stages of diversification and the genetic consequences of postglacial migration of a hermaphroditic polyploid plant, Cerastium alpinum. It has a continuous distribution in the alpine region, where it grows on alpine heaths and serpentine soils that are rich in heavy metals. Within the boreal forest C. alpinum has a scattered distribution on serpentine, dolomite and steep slopes. Plants from 31 populations in Norway, Sweden and Finland were subjected to enzyme electrophoresis. Analyses of the enzyme phenotypes suggest that C. alpinum has colonized Fennoscandia through two postglacial immigration events resulting in a southeastern and a southwestern lineage. These two lineages seem to meet in a hybrid zone in northern Sweden. Large genetic differences were found among most populations in both the southeastern and the southwestern lineages. This suggests that the populations are effectively isolated from each other.

We analyzed the genetic basis of morphological differences between two wild species of teosinte (Zea diploperennis and Z. mays ssp. parviglumis), which are relatives of maize. These two species differ in a number of taxonomically important traits including the structure of the tassel (male inflorescence), which is the focus of this report. To investigate the genetic inheritance of six tassel traits, quantitative trait locus (QTL) mapping with 95 RFLP markers was employed on a population of 425 F2 plants. Each trait was analyzed by interval mapping (IM) and composite interval mapping (CIM) to identify and characterize the QTL controlling the differences in tassel mor- phology. We detected two to eight QTL for each trait. In total, 30 QTL with IM and 33 QTL with CIM were found for tassel morphology. QTL for several of the traits mapped near each other, suggesting pleiotropy and/or linkage of QTL. The QTL showed small to moderate magnitudes of effect. No QTL of exceptionally large effect were found as seen under domestication and in the case of some other natural species. Thus, the model involving major QTL of large effect seems not to apply to the traits and species analyzed. A mixture of QTL with positive and negative allelic effects was found for most tassel traits and may suggest a history of periodic changes in the direction of selection during the divergence of Z. diploperennis and Z. mays ssp. parviglumis or fixation of QTL alleles by random genetic drift rather than selection.

Silene dioica (Caryophyllaceae) has evolved different morphotypes that vary in hairiness in different habitats in the Swedish mountains. A hairy form, var. serpentinicola and a hairless (glabrous) form, var. smithii grow on serpentine, i.e. a dry soil rich in heavy metals. Var. lapponica is a densely hairy morphotype on subalpine meadows. Crosses between these varieties showed that glabrousness has a simple Mendelian inheritance and that glabrousness is determined by a recessive allele. The crosses further suggested that the recessive allele has been present in the original population that invaded the mountain area. Glabrous plants have, however, never been found on subalpine meadows, even though an earlier study has shown that gene flow is relatively common among serpentine and nearby meadow populations. In addition to the genetic background of glabrousness, we studied the grazing on glabrous smithii and hairy serpentinicola and lapponica plants by the slugs Arion fasciatus and Arion subfuscus and the snail Arianta arbustorum in greenhouse and in nature. Arion slugs, abundant on subalpine meadows, preferred glabrous plants and, in particular, A. fasciatus consumed high amounts of S. dioica. Arianta snails, however, consumed irrespective of morphotype. The selective behaviour was seen both at the seedling stage and the adult plant stage. Arion slugs were more choosy when they were given a choice of glabrous smithii and densely hairy lapponica plants then when they were served smithii and the less hairy variety serpentinicola. This study implies that the Arion slugs are important agents of selection on subalpine meadows. The selective grazing seems to counteract the flow of the recessive allele that determines glabrousness, from serpentine into meadow populations. Gastropods are rare in the dry serpentine habitat. The evolution of a distinct glabrous morphotype on serpentine is therefore rather due to relaxation of selection than to adaptation to serpentine stress.

Silene dioica (L.) Clairv. (Caryophyllaceae) is an insect pollinated and obligately cross-fertilising herb commonly found on serpentine, i.e. soils rich in heavy metals, and on cultivated meadows. Groups of Silene populations were studied in two areas in the Scandinavian mountains. In a northern area Silene populations grow in an open pine and birch forest while in the south, populations are surrounded by a continuous and dense spruce forest. Gene flow, estimated on the basis of allele frequencies, is highly restricted in the south in comparison to the north. This has led to an extensive genetic differentiation among populations. The pollinator guilds (Thricops flies, syrphid flies and bumblebees) are similar in the northern and southern populations and can therefore not account for the difference in gene flow. The flowering in nearby populations overlaps broadly and is also not the reason for the genetic differentiation in the south. The differentiation is more likely due to vegetation that limits pollinator movement. The level of gene flow differs in the two habitats both in the north and in the south. The gene flow is restricted among serpentine populations but common among meadow populations. Pollen dispersal, and in particular, anthropogenic seed dispersal mediate gene flow among meadows. Serpentine populations are unaffected by human activity. Pollinators are therefore the main agent of gene flow on serpentine. This study shows the importance of the surrounding vegetation in influencing gene flow patterns among populations as well as that habitat fragmentation affects the genetic properties of plant populations. In addition, we have also observed a restricted gene flow within populations. Enzyme allele frequencies show an excess of homozygotes within most of the populations. This can be explained through limited pollen dispersal and differences in male and female flowering density.

Serpentine soils are rich in heavy metals and have a distinctive flora. Silene dioica is a member of the Scandinavian serpentine plant community but is also widespread outside serpentine soils. To study the population genetic consequences of serpentine stress and the origin and evolution of serpentine populations we analyzed the isozyme genetic structure of S. dioica. Seventeen pop- ulations located in the mountains of Vaisterbotten and Jamtland, central Sweden, were investigated by starch gel enzyme electrophoresis. About one half of the populations grow in serpentine soils and the rest on adjacent non-serpentine sites. Analyses of allele frequencies show that both ser- pentine and non-serpentine populations in the northern part of the studied area (Vaisterbotten) are genetically similar. Evidently serpentine does not exert strong selection acting upon isozyme loci. In the south (Jaimtland), however, the serpentine populations exhibit genetic differentiation. This allozyme divergence is probably not due to direct selection but rather represents the effects of isolation and genetic drift. The results suggest that S. dioica has colonized serpentine repeatedly and that the tolerant populations have a multiple origin.