浑善达克沙地几种优势植物的生态适应

浑善达克沙地是我国北方干旱半干旱区的主要沙地和严重荒漠化地区之一。同时也是我国内陆沙尘的主要源区之一。研究当地植物对沙地环境的适应方式有助于当地的荒漠化治理，加快生态环境恢复。本文应用（温室和野外）实验生态学方法研究了沙地植物对水分、沙埋和养分异质性环境的生态适应对策。 浑善达克沙地属于半干旱区域。在一次降雨过后，适合植物生长的湿沙层逐渐下降，或者说不适合植物生长的干沙层逐渐加厚。为了能够有效地从沙层中获得水分，植物的根系的伸长的速度必须大于湿沙层下降的速度。因此，当地分布的植物的根系则应该具有相应的形态、生长和生物量投资方面的可塑性。羊柴（Hedysarum laeve Maxim. (Leguminosae) ）是当地的主要半灌木和固沙植物。我们研究了羊柴幼苗对不同土壤含水量和不同模拟降雨量的反应格局，以及移动沙丘生长季节的土壤含水量变动规律。发现，大于3％土壤含水量适合羊柴幼苗生长。最适土壤含水量为12%－20% 。在生长季节早期，羊柴幼苗比较容易在沙丘上定居。如果没有充分的后续降雨的话，萌发后的幼苗在沙丘顶部、落沙坡中部和后部定居比较困难。降雨量越大，根系越深、总根长越大、根系直径越大、侧根越多、根冠比越低。 在干旱半干旱区域的沙地，种子萌发的时间和地点对于植物的定居和存活具有关键作用。通过温室实验，研究了小叶锦鸡儿和羊柴的种子萌发和出苗对不同深度的沙埋的反应，及其与种子大小的关联。结果表明，小叶锦鸡儿和羊柴的种子萌发和出苗对浑善达克沙地的自然生境表现出相似的生态适应。这两个种的种子萌发的适宜温度为10-15ºC, 这是当地春季的平均温度。由于这两个种的种子萌发是光敏感的，所以它们的种子萌发受土壤含水量和光照强度二者之间的平衡调节。2 cm 的较浅沙埋最有利于种子萌发和出苗。种子埋得越深，种子的萌发率和出苗率越低，更多的种子受胁迫休眠保存在土壤中作为种子库。不同大小的种子的不同萌发能力也是一种重要的生态适应，这可以调节种子在合适的时间和合适的沙埋深度萌发和出苗。 冰草（Agropyron cristatum (L.) Gaertn）是浑善达克沙地植物群落中占优势的多年生根茎草本植物之一。种子在合适的地点和时间萌发以及幼苗在合适的地点和时间生长，对于冰草在沙地环境条件下生存和分布具有重要意义。本文研究了浑善达克沙地4-10月份土壤含水量变动情况和冰草种子萌发、出苗和幼苗生长对土壤含水量的响应。结果表明，4月下旬至5月上中旬的土壤含水量对冰草种子萌发、出苗和定居极为关键。控制条件下，冰草种子萌发的最适土壤含水量是12%-20%，出苗以及幼苗生长的最适土壤含水量是12%-16%。当土壤含水量低于3%，冰草种子不能萌发，土壤含水量低于6%时，幼苗不能出土和定居。当土壤含水量达到16%时，冰草幼苗生物量有所下降。在6%-8%的土壤含水量条件下，植株将更多的生物量投资于根的生长。 本文对浑善达克沙地低湿滩地、滩地-风沙沉积过渡区到风沙沉积区的赖草（Leymus secalinus （Georgi） Tzvel.）分株数、地上生物量、土壤水分和养分异质性进行了研究。结果表明，从低湿滩地、滩地-风沙沉积过渡区到风沙沉积区，随着土壤养分的降低，赖草分株数和地上生物量反而增加。赖草的分株数在三种生境中都存在空间自相关。并且，从低湿滩地、过渡区到风沙沉积区空间自相关的变程逐渐增加。地上生物量同样存在空间自相关，但变程以过渡区最大，风沙沉积区最低。土壤水分仅仅在风沙沉积区存在空间自相关。在三种生境条件下，土壤全氮和有机质的空间分布格局相似，都在低湿滩地和过渡区存在空间自相关且变程相似。在风沙沉积区不存在空间自相关。赖草的空间分布格局在低湿滩地为偏离随机适度聚集的分布格局，在过渡区近似于聚居分布，而在风沙沉积区为均匀分布格局。 豆科锦鸡儿属（Caragana ）植物因其可以生物固氮而在草原生态系统当中具有特殊的地位。定量分析小叶锦鸡儿（Caragana microphylla ）灌丛在浑善达克沙地不同生境条件下的分布格局，有助于理解植被与土壤养分循环之间的关系和合理制定退化沙地的恢复对策。本文采用地统计学中的半方差分析和分形分析两种方法研究了浑善达克沙地小叶锦鸡儿灌丛的空间分布格局，分析了不同生境中小叶锦鸡儿灌丛的植被、土壤水分和养分间的相关性。结果表明（1）小叶锦鸡儿灌丛在三种生境条件下（滩地、固定沙丘和半固定沙丘）的盖度、土壤有机质和全氮的空间分布符合球状模型，空间自相关显著;（2）土壤pH值在滩地和固定沙丘的空间分布符合球状模型且空间自相关显著，但在半固定沙丘空间自相关不显著;（3）土壤水分在三种生境中的空间分布都符合线性模型，空间自相关显著。（4）植被的变异尺度（变程）小于各个土壤要素的变异尺度。植被（小叶锦鸡儿灌丛）的分布和形成过程决定了土壤养分的分布和形成过程。当地的豆科植物在养分“沃岛”现象的形成中起了重要作用，合理的利用和布局当地的豆科植物，可以更加有效地补充草地生态系统的养分，从而有利于加快当地生态系统的恢复进程。

Phenotypic changes in artemisinin-resistant plasmodium falciparum lines in vitro: evidence for decreased sensitivity to dormancy and growth inhibition

The appearance of Plasmodium falciparum parasites with decreased in vivo sensitivity but no measurable in vitro resistance to artemisinin has raised the urgent need to characterize the artemisinin resistance phenotype. Changes in the temporary growth arrest (dormancy) profile of parasites may be one aspect of this phenotype. In this study, we investigated the link between dormancy and resistance, using artelinic acid (AL)-resistant parasites. Our results demonstrate that the AL resistance phenotype has (i) decreased sensitivity of mature-stage parasites, (ii) decreased sensitivity of the ring stage to the induction of dormancy, and (iii) a faster recovery from dormancy.

Regulation of dormancy in barley by blue light and after-ripening : effects on abscisic acid and gibberellin metabolism

White light strongly promotes dormancy in freshly harvested cereal grains, whereas dark and after-ripening have the opposite effect. We have analyzed the interaction of light and after-ripening on abscisic acid (ABA) and gibberellin (GA) metabolism genes and dormancy in barley (Hordeum vulgare ‘Betzes’). Analysis of gene expression in imbibed barley grains shows that different ABA metabolism genes are targeted by white light and after-ripening. Of the genes examined, white light promotes the expression of an ABA biosynthetic gene, HvNCED1, in embryos. Consistent with this result, enzyme-linked immunosorbent assays show that dormant grains imbibed under white light have higher embryo ABA content than grains imbibed in the dark. After-ripening has no effect on expression of ABA biosynthesis genes, but promotes expression of an ABA catabolism gene (HvABA8′OH1), a GA biosynthetic gene (HvGA3ox2), and a GA catabolic gene (HvGA2ox3) following imbibition. Blue light mimics the effects of white light on germination, ABA levels, and expression of GA and ABA metabolism genes. Red and far-red light have no effect on germination, ABA levels, or HvNCED1. RNA interference experiments in transgenic barley plants support a role of HvABA8′OH1 in dormancy release. Reduced HvABA8′OH1 expression in transgenic HvABA8′OH1 RNAi grains results in higher levels of ABA and increased dormancy compared to nontransgenic grains.

Conservation and divergence of four kiwifruit SVP-like MADS-box genes suggest distinct roles in kiwifruit bud dormancy and flowering

MADS-box genes similar to Arabidopsis SHORT VEGETATIVE PHASE (SVP) have been implicated in the regulation of flowering in annual species and bud dormancy in perennial species. Kiwifruit (Actinidia spp.) are woody perennial vines where bud dormancy and out-growth affect flower development. To determine the role of SVP-like genes in dormancy and flowering of kiwifruit, four MADS-box genes with homology to Arabidopsis SVP, designated SVP1, SVP2, SVP3, and SVP4, have been identified and analysed in kiwifruit and functionally characterized in Arabidopsis. Phylogenetic analysis indicate that these genes fall into different sub-clades within the SVP-like gene group, suggesting distinct functions. Expression was generally confined to vegetative tissues, and increased transcript accumulation in shoot buds over the winter period suggests a role for these genes in bud dormancy. Down-regulation before flower differentiation indicate possible roles as floral repressors. Over-expression and complementation studies in Arabidopsis resulted in a range of floral reversion phenotypes arising from interactions with Arabidopsis MADS-box proteins, but only SVP1 and SVP3 were able to complement the svp mutant. These results suggest that the kiwifruit SVP-like genes may have distinct roles during bud dormancy and flowering.

A rapid transcriptional activation is induced by the dormancy-breaking chemical hydrogen cyanamide in kiwifruit (Actinidia deliciosa) buds

Budbreak in kiwifruit (Actinidia deliciosa) can be poor in locations that have warm winters with insufficient winter chilling. Kiwifruit vines are often treated with the dormancy-breaking chemical hydrogen cyanamide (HC) to increase and synchronize budbreak. This treatment also offers a tool to understand the processes involved in budbreak. A genomics approach is presented here to increase our understanding of budbreak in kiwifruit. Most genes identified following HC application appear to be associated with responses to stress, but a number of genes appear to be associated with the reactivation of growth. Three patterns of gene expression were identified: Profile 1, an HC-induced transient activation; Profile 2, an HC-induced transient activation followed by a growth-related activation; and Profile 3, HC- and growth-repressed. One group of genes that was rapidly up-regulated in response to HC was the glutathione S-transferase (GST) class of genes, which have been associated with stress and signalling. Previous budbreak studies, in three other species, also report up-regulated GST expression. Phylogenetic analysis of these GSTs showed that they clustered into two sub-clades, suggesting a strong correlation between their expression and budbreak across species.

Overexpression of the kiwifruit SVP3 gene affects reproductive development and suppresses anthocyanin biosynthesis in petals, but has no effect on vegetative growth, dormancy, or flowering time

SVP-like MADS domain transcription factors have been shown to regulate flowering time and both inflorescence and flower development in annual plants, while having effects on growth cessation and terminal bud formation in perennial species. Previously, four SVP genes were described in woody perennial vine kiwifruit (Actinidia spp.), with possible distinct roles in bud dormancy and flowering. Kiwifruit SVP3 transcript was confined to vegetative tissues and acted as a repressor of flowering as it was able to rescue the Arabidopsis svp41 mutant. To characterize kiwifruit SVP3 further, ectopic expression in kiwifruit species was performed. Ectopic expression of SVP3 in A. deliciosa did not affect general plant growth or the duration of endodormancy. Ectopic expression of SVP3 in A. eriantha also resulted in plants with normal vegetative growth, bud break, and flowering time. However, significantly prolonged and abnormal flower, fruit, and seed development were observed, arising from SVP3 interactions with kiwifruit floral homeotic MADS-domain proteins. Petal pigmentation was reduced as a result of SVP3-mediated interference with transcription of the kiwifruit flower tissue-specific R2R3 MYB regulator, MYB110a, and the gene encoding the key anthocyanin biosynthetic step, F3GT1. Constitutive expression of SVP3 had a similar impact on reproductive development in transgenic tobacco. The flowering time was not affected in day-neutral and photoperiod-responsive Nicotiana tabacum cultivars, but anthesis and seed germination were significantly delayed. The accumulation of anthocyanin in petals was reduced and the same underlying mechanism of R2R3 MYB NtAN2 transcript reduction was demonstrated.

Artemisinin resistance in Plasmodium falciparum : a process linked to dormancy?

Artemisinin (ART) based combination therapy (ACT) is used as the first line treatment of uncomplicated falciparum malaria in over 100 countries and is the cornerstone of malaria control and elimination programs in these areas. However, despite the high potency and rapid parasite killing action of ART derivatives there is a high rate of recrudescence associated with ART monotherapy and recrudescence is not uncommon even when ACT is used. Compounding this problem are reports that some parasites in Cambodia, a known foci of drug resistance, have decreased in vivo sensitivity to ART. This raises serious concerns for the development of ART resistance in the field even though no major phenotypic and genotypic changes have yet been identified in these parasites. In this article we review available data on the characteristics of ART, its effects on Plasmodium falciparum parasites and present a hypothesis to explain the high rate of recrudescence associated with this potent class of drugs and the current enigma surrounding ART resistance.

Artemisinin-induced parasite dormancy : a plausible mechanism for treatment failure

Background Artemisinin-combination therapy is a highly effective treatment for uncomplicated falciparum malaria but parasite recrudescence has been commonly reported following artemisinin (ART) monotherapy. The dormancy recovery hypothesis has been proposed to explain this phenomenon, which is different from the slower parasite clearance times reported as the first evidence of the development of ART resistance. Methods In this study, an existing P. falciparum infection model is modified to incorporate the hypothesis of dormancy. Published in vitro data describing the characteristics of dormant parasites is used to explore whether dormancy alone could be responsible for the high recrudescence rates observed in field studies using monotherapy. Several treatment regimens and dormancy rates were simulated to investigate the rate of clinical and parasitological failure following treatment. Results The model output indicates that following a single treatment with ART parasitological and clinical failures occur in up to 77% and 67% of simulations, respectively. These rates rapidly decline with repeated treatment and are sensitive to the assumed dormancy rate. The simulated parasitological and clinical treatment failure rates after 3 and 7 days of treatment are comparable to those reported from several field trials. Conclusions Although further studies are required to confirm dormancy in vivo, this theoretical study adds support for the hypothesis, highlighting the potential role of this parasite sub-population in treatment failure following monotherapy and reinforcing the importance of using ART in combination with other anti-malarials.

Artemisinin induced dormancy in Plasmodium falciparum : duration, recovery rates and implications in treatment failure

Background Despite the remarkable activity of artemisinin and its derivatives, monotherapy with these agents has been associated with high rates of recrudescence. The temporary arrest of the growth of ring-stage parasites (dormancy) after exposure to artemisinin drugs provides a plausible explanation for this phenomenon. Methods Ring-stage parasites of several Plasmodium falciparum lines were exposed to different doses of dihydroartemisinin (DHA) alone or in combination with mefloquine. For each regime, the proportion of recovering parasites was determined daily for 20 days. Results Parasite development was abruptly arrested after a single exposure to DHA, with some parasites being dormant for up to 20 days. Approximately 50% of dormant parasites recovered to resume growth within the first 9 days. The overall proportion of parasites recovering was dose dependent, with recovery rates ranging from 0.044% to 1.313%. Repeated treatment with DHA or with DHA in combination with mefloquine led to a delay in recovery and an ∼10-fold reduction in total recovery. Strains with different genetic backgrounds appeared to vary in their capacity to recover. Conclusions These results imply that artemisinin-induced arrest of growth occurs readily in laboratory-treated parasites and may be a key factor in P. falciparum malaria treatment failure.

Grain dormancy QTL identified in a doubled haploid barley population derived from two non-dormant parents

Grain dormancy provides protection against pre-harvest sprouting (PHS) in cereals. Composite interval mapping and association analyses were performed to identify quantitative trait loci (QTL) contributing grain dormancy in a doubled haploid (DH) barley population (ND24260 x Flagship) consisting of 321 lines genotyped with DArT markers. Harvest-ripe grain collected from three field experiments was germinated over a 7-day period to determine a weighted germination index for each line. DH lines displaying moderate to high levels of grain dormancy were identified; however, both parental lines were non-dormant and displayed rapid germination within the first two days of testing. Genetic analysis identified two QTL on chromosome 5H that were expressed consistently in each of the three environments. One QTL (donated by Flagship) was located close to the centromeric region of chromosome 5H (qSDFlag), accounting for up to 15% of the phenotypic variation. A second QTL with a larger effect (from ND24260) was detected on chromosome 5HL (qSDND), accounting for up to 35% of the phenotypic variation. qSDFlag and qSDND displayed an epistatic interaction and DH lines that had the highest levels of grain dormancy carried both genes. We demonstrate that qSDND in the ND24260 9 Flagship DH population is positioned proximal and independent to the well-characterised SD2 region that is associated with both high levels of dormancy and inferior malt quality. This indicates that it should be possible to develop cultivars that combine acceptable malting quality and adequate levels of grain dormancy for protection against PHS by utilizing these alternate QTL.

Fine Mapping of qDor7, a Major QTL Affecting Seed Dormancy in Sorghum (Sorghum bicolor (L.) Moench)

Seed dormancy is a key domestication trait for major crops, which is acquired in long-term systems development processes and enables the survival of plants in adverse natural conditions. It is a complex trait under polygenic control and is affected by endogenous and environmental factors. In the present study, a major seed dormancy QTL in sorghum (Sorghum bicolor (L.) Moench), qDor7, detected previously, was fine mapped using a large, multi-generational population. The qDor7 locus was delimited to a 96-kb region which contains 16 predicted gene models. These results lay a solid foundation for cloning qDor7. In addition, the functional markers tightly linked to the seed dormancy QTL may be used in marker-assisted selection for seed dormancy in sorghum.

Time to rest : Signals in shoot apex developmental transitions underlying dormancy

Vuodenajat rytmittävät monivuotisten kasvien elämää pohjoisella pallonpuoliskolla, jolla varmin merkki lähestyvästä talvikaudesta on asteittain lyhenevä päivänpituus. Kun päivänpituus on lyhentynyt tiettyyn raja-arvoon saakka, kasvu hiipuu ja kasvin kehityksessä tapahtuu suuria muutoksia. Väitöskirjatyössäni tutkittiin mekanismeja, jotka liittyvät pituuskasvun päättymiseen, silmujen lepotilan kehittymiseen ja kärkisilmun muodostumiseen hybridihaavan ja koivuntaimilla lyhyen päivänpituuden seurauksena kasvihuoneolosuhteissa. Vain lepotilaiset silmut selviytyvät luonnossa ankaran talvikauden yli, joten etenkin lepotilan kehittymisen tutkiminen on keskeistä pyrittäessä selvittämään monivuotisille kasveille tyypillisen kasvutavan mekanismeja. Jo pitkään on tiedetty, että täysikasvuiset lehdet vastaanottavat tiedon päivänpituudesta ja lähettävät signaaleja varren johtojänteissä ylöspäin kohti kasvin kärkiosaa. Sen sijaan varren kärjen ja kärkikasvupisteen roolia lepotilan kehittymisessä on selvitetty vain vähän. Kuitenkin juuri kärkikasvupisteen selviytyminen vuodesta toiseen on tärkeää, koska sen jakautumiskykyiset solukot tuottavat kasvin maanpäälliset osat. Tässä työssä tehdyissä varttamiskokeissa osoitettiin, että varren kärki ei ainoastaan vastaanota signaaleja lehdistä ja ajoita toimintaansa niiden mukaan, vaan myös kärjellä itsellään on aktiivinen rooli lepotilan kehittymisessä. Erityisesti kiinnitettiin huomiota kärkikasvupisteen eri alueiden, ns. apikaalimeristeemin ja rib-meristeemin erilaisiin tehtäviin ja pääteltiin, että molemmat vaikuttavat lepotilan kehittymiseen. Kokeissa käytettiin normaalien hybridihaapojen lisäksi siirtogeenisiä hybridihaapoja, jotka eivät lopeta kasvuaan lyhyt päivä –olosuhteissa. Siirtogeeniset hybridihaavat ilmensivät voimakkaasti fytokromi A -nimistä valon vastaanottajamolekyyliä rib-meristeemin alueella, mikä saattoi osaltaan vaikuttaa poikkeavaan pituuskasvukäyttäytymiseen. Myös useiden lepotilan kehittymiseen liittyvien geenien ilmenemisessä havaittiin poikkeavuuksia verrattuna ei-siirtogeenisiin kontrolleihin, joiden silmuissa kehittyi lepotila lyhyt päivä –altistuksen seurauksena. Väitöskirjatyössäni havaittiin, että myös kaasumainen kasvihormoni etyleeni toimii viestinvälittäjänä silmujen lepotilan kehittymisessä ja vaikuttaa etenkin lepotilan oikeaan ajoittumiseen. Etyleenillä huomattiin olevan määräävä rooli päätesilmun muodostumisessa: siirtogeeniset koivut, jotka eivät aisti etyleeniä, eivät muodostaneet päätesilmua. Silti siirtogeeniset koivut vaipuivat lepotilaan, joskin myöhemmin kuin ei-siirtogeeniset kontrollit. Tämän perusteella todettiin, että lepotilan ja päätesilmun kehittyminen ovat erillisiä kehitystapahtumia, vaikka ne saattavatkin ajoittua osaksi päällekkäin.

Chelation-enforced metal-arene interactions : insights into substrate binding and catalysis by late transition metal complexes

Understanding and catalyzing chemical reactions requiring multiple electron transfers is an endeavor relevant to many outstanding challenges in the field of chemistry. To study multi-electron reactions, a terphenyl diphosphine framework was designed to support one or more metals in multiple redox states via stabilizing interactions with the central arene of the terphenyl backbone. A variety of unusual compounds and reactions and their relevance toward prominent research efforts in chemistry are the subject of this dissertation.

Chapter 2 introduces the para-terphenyl diphosphine framework and its coordination chemistry with group 10 transition metal centers. Both mononuclear and dinuclear compounds are characterized. In many cases, the metal center(s) are stabilized by the terphenyl central arene. These metal–arene interactions are characterized both statically, in the solid state, and fluxionally, in solution. As a proof-of-principle, a dinickel framework is shown to span multiple redox states, showing that multielectron chemistry can be supported by the coordinatively flexible terphenyl diphosphine.

Chapter 3 presents reactivity of the terphenyl diphosphine when bound to a metal center. Because of the dearomatizing effect of the metal center, the central arene of the ligand is susceptible to reactions that do not normally affect arenes. In particular, Ni-to-arene H-transfer and arene dihydrogenation reactions are presented. Additionally, evidence for reversibility of the Ni-to-arene H-transfer is discussed.

Chapter 4 expands beyond the chelated metal-arene interactions of the previous chapters. A dipalladium(I) terphenyl diphosphine framework is used to bind a variety of exogenous organic ligands including arenes, dienes, heteroarenes, thioethers, and anionic ligands. The compounds are structurally characterized, and many ligands exhibit unprecedented bindng modes across two metal centers. The relative binding affinities are evaluated spectroscopically, and equilibrium binding constants for the examined ligands are determined to span over 13 orders of magnitude. As an application of this framework, mild hydrogenation conditions of bound thiophene are presented.

Chapter 5 studies nickel-mediated C–O bond cleavage of aryl alkyl ethers, a transformation with emerging applications in fields such as lignin biofuels and organic methodology. Other group members have shown the mechanism of C–O bond cleavage of an aryl methyl ether incorporated into a meta-terphenyl diphosphine framework to proceed through β-H elimination of an alkoxide. First, the electronic selectivity of the model system is examined computationally and compared with catalytic systems. The lessons learned from the model system are then applied to isotopic labeling studies for catalytic aryl alkyl ether cleavage under dihydrogen. Results from selective deuteration experiments and mass spectrometry draw a clear analogy between the mechanisms of the model and catalytic systems that does not require dihydrogen for C–O bond cleavage, although dihydrogen is proposed to play a role in catalyst activation and catalytic turnover.

Appendix A presents initial efforts toward heterodinuclear complexes as models for CO dehydrogenase and Fischer Tropsch chemistry. A catechol-incorporating terphenyl diphosphine is reported, and metal complexes thereof are discussed.

Appendix B highlights some structurally characterized terphenyl diphosphine complexes that either do not thematically belong in the research chapters or proved to be difficult to reproduce. These compounds show unusual coordination modes of the terphenyl diphosphine from which other researchers may glean insights.

Experimental research on dormancy in Cyclops vicinus [Translation from: Naturwissenschaften 56 93-94, 1969]

Dormancy has been observed in several species of Cyclops. Animals in dormancy are characterised by almost complete inactivity of the main appendages and sluggish intestinal peristalsis. In Cyclops vicinus dormancy can take place in copepod stages III (19%), IV (78%) and V (3%) but one and the same animal only in one developmental stage. The author gives his own results between frequency of dormancy and photoperiod (20 C, 1000 lux). He concludes that dormancy in C. vicinus can be influenced by day length and can be ended prematurely by short day length.