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资助项目
6 could use lots of photosynthates, but contributed little to the accumulation of biomass. 4. Photosynthetic rate of P. armeniacum decreased a little at the noon, and the highest photosynthetic rate was observed at 10:00h in the greenhouse. The variation of photosynthetic rate was in the same trend as stomatal conductance. Higher relative humidity seemed to be the key for higher photosynthetic rate in P. armeniacum. 5. The photosynthetic capacity of C. flavum was statistically larger than that of P. armeniacum. The lower leaf photosynthetic capacity of P. armeniacum was related to its lower leaf nitrogen concentration,leaf phosphorus concentration and enzyme activities. Meanwhile, the extremely lower stomatal conductance and internal mesophyll conductance might greatly limit the photosynthetic capacity of P. armeniacum. The lower stomatal conductance and photosynthetic rate of Paphiopedilum might partially caused by the lack of chloroplasts in the guard cell of Paphiopedilum. Compared with C. flavum, P. armeniacum was more fond of shade environment.6. The short longevity leaf of Cypripedium had bigger photosynthetic capacity and greater potential for fast growth. But the longer LL of Paphiopedilum enhanced nutrient conservation which could compensate its lower photosynthetic capacity. The short longevity leaf of Cypripedium usually had higher photosynthetic rate per unit leaf mass and dark respiration rate, and photosynthetic capacity decreased fast with leaf age. However, for Paphiopedilum, the situation was the opposite. 7. Compared with Cypripedium, Paphiopedilum had higher water use efficiency and lower photosynthetic nitrogen use efficiency. 8. The leaf of Paphiopedilum had higher leaf construction cost and longer repayment time than that of Cypripedium. The leaf structures and physiological functions of Paphiopedilum and Cypripedium reflected the adaptation to their habitats. The leaf morphological and physiological evolution of Paphiopedilum was related to water and resource-conserving traits in the karst habitat. The leaf traits of Cypripedium were the adaptation to the environment rich in water and nutrients but easy to change with seasons.Our results provided evidence of divergent evolution of congeneric orchids under natural 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and Cypripedium，known as slipper orchids in horticulture, belong to the subfamily Cypripedioideae of the Orchidaceae. Although they are closely related phylogenetically, there are significant differences in leaf traits and geographical distributions between two genera. This dissertation includes the following sections: (1) the leaf functional traits were compared in six species of the two genera; (2) the physiological responses of P. armeniacum to different water regimes, light regimes and low temperature; (3) the leaf phenotypic plastics of C. flavum in response to the different light condition and the photosynthetic characteristics of three Cypripedium species during sexual reproduction. The aims are to understand the convergent and divergent evolution between the two genera in leaf traits and their adaptive significances, and the leaf plastic responses to different levels of resources. Such information could provide scientific basis for conservation and domestication of Paphiopedilum and Cypripedium. The results are given below:1. Compared with Paphiopedilum, Cypripedium showed significantly higher photosynthetic rate (Pmax), leaf nitrogen content (Na), photosynthetic nitrogen utilization (PNUE), the fractions of leaf nitrogen partitioning in carboxylation (PC) and bioenergetics (PB), specific leaf area (SLA), ratio of leaf chlorophyll a and b (Chla/b), but significantly lower leaf construction cost (CC) and the ratio of leaf carbon content to leaf nitrogen (C/N). These leaf traits of Cypripedium are considered as the adaptation to short growing period and rich soil nutrients in the alpine habitats. Conversely, the long life span, low Pmax and mesophyll conductance (gm) but high SLA, CC and C/N in Paphiopedilum indicated that the adaptation to low-light, limited-nutrient habitat in the limestone area. As a sympatric species of Paphiopedilum, C. lentiginosum not only kept phylogenetically leaf traits of Cypripedium, suchas stomatal conductance (gs), Pmax, PNUE and dormant in winter, but also possessed many leaf traits which is similar to that in Paphiopedilum, such as relative stomatal limitations (RSL), gm, the ratio of leaf chlorophyll a and b (Chl a/b), fraction of leaf nitrogen allocated to light-harvesting components (PL). These results indicated the convergent and divergent evolution of Paphiopedilum and Cypripedium in leaf traits.2. Paphiopedilum. armeniacum exhibited a high plasticity of leaf photosynthetic function in response to different light regimes, but the responses changes with the time. Due to grow under low light habitat, P. armeniacum grown under 50% shade (HL) had the significantly lowest Pmax than the plants grown under 75% shade (ML) and 95% shade (LL) after six months. However, after twelve months, the Pmax of the plants grown under HL increased significantly and then became the highest one among three levels of light. It is also found that leaf dry mass per unit area (LMA), leaf stomatal conductance (gS), internal mesophyll conductance (gm), the fraction of leaf nitrogen partitioning in photosynthetic carboxylation (PC), bioeneretics (PB) were greatly influenced by irradiance. The plants grown under HL increased gS, gm, PC, PB to increase Pmax. In addition, the plants grown under HL had the highest ratio of total chlorophyll content to total Carotenoid content (Car/Chl) while the plants grown under LL had the lowest ratio of leaf chlorophyll a and b (Chl a/b). As a result, plasticity of leaf photosynthetic physiology of P. armeniacum in response to different light regimes depended largely on leaf nitrogen partitioning and leaf structure. As for the numbers of flowering and fruiting, ML was the best light level.3. The responses of P. armeniacum to different water regimes were not significantly different. But the Pmax and the maximum photochemical efficiency of PSⅡ (Fv/Fm) decreased with the increased frequency of watering. The reasons were that the plants have high respiration rate (Rd) and make more use of light energy to oxidation cycle. The plants watered every eight days (MW) and every twenty days (LW) had higher Pmax than the plant watered every four days (HW) mainly because of the higher PC and PB. Besides, the leaves of P. armeniacum had excellent property for holding water also contributed to the high photosynthetic capacity.4. Paphiopedilum. armeniacum was very sensitive to the low temperature. The plants significantly decreased photosynthetic capacity after grown under 4℃ for three days and the photosynthetic machinery was destroyed after fifteen days. The photosynthetic capacity of P. armeniacum exhibited no change at 10℃ and 15℃.5. Cypripedium flavum of four habitats (DB, XRD, XZD and TSQ) with different light intensity exhibited different photosynthetic characteristics after transplanted to the same environment in Kunming. Among the habitats, the light intensity of DB was the highest while XRD was the lowest. The light intensity of XZD and TSQ were not significantly difference. Among all the plants in Kunming, the plants of DB had the significantly highest Pmax but the plants of XRD had the lowest Pmax. The light saturation point (LSP) and photosynthetic nitrogen use efficiency (PNUE) agreed well with the light intensity of four habitats and contributed to the high Pmax of DB. The LMA, Chl and leaf nitrogen content were not different among all the plants. C. flavum exhibited sensitively response to the change of light in leaf construction while kept the plasticity of leaf photosynthetic characteristics which developed from its own habitat.6. The photosynthetic capacity of C. tibeticum and C. flavum were significantly increased at the flowering stage. For these two species, the significantly increased Amax were closely related to the maximum carboxylation rate by ribulose-1, 5-bisphosphate carboxylase/oxygenase (Vcmax), photon saturated rate of electron transport (Jmax), the rate of triose phosphate utilization (TPU) and actual quantum efficiency of the photosystem II photochemistry (ΦPSII) respectively. However, flowering almost did not affect the photosynthetic capacity of C. guttatum. C. guttatum had the smallest plant size, the leaf area, the volume of labellum and the volume of fruit, but the biggest fruit volume per leaf area among three species. These results indicated that for C. flavum and C. tibeticum there were a physiological mechanism in photosynthesis to compensate the cost of flowering as well as increased resource acquisitions, which would be beneficial to the survival or future flowering of the plant. C. gutattum could keep a steady photosynthetic capacity during life history. This kind of pattern could decrease the effect of the reproductive costs as much as possible. In contrast to C. flavum and C. tibeticum, C. gutattum possessed a more economical and effective reproductive pattern which maybe related to its wider distribution.In conclusion, Paphiopedilum and Cypripedium have significantly different leaf traits which agree well with their habitats and there is a divergent and convergent evolution between the two genera. P. armeniacum is much tolerant and responsive to varying water and light availability but very sensitivity to the low temperature. Confronting the suddenly change of light environment, C. flavum can respond sensitively to the change of light in leaf construction but the plasticity of leaf photosynthetic characteristics which developed from its own habitat can hold for the next growing season. In contrast to C. flavum and C. tibeticum, C. gutattum possesses a more economical and effective reproductive pattern which maybe related to its wider distribution. The study of the relationship between the two genera, the response and tolerance to the environmental factors of the two genera are important for understanding the adaptation and evolution of the 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sect. Cyathophora is a distinctive group endemic to the eastern Himalaya-Hengduan Mountains region. It was regarded as a ‘grex’ or section and included all four general corolla types of Pedicularis. A unique feature is that the leaf and bract bases are fused together to form a cup-like structure around the stem at each node. Our molecular phylogenetic study indicated that P. sect. Cyathophora was a monophyletic lineage, although the relationships among species were not fully resolved. P. rex C. B. Clarke ex Maxim. is the most various and wide-distributed species in P. sect. Cyathophora. It is extremely variable in wild populations, and elicits a vast quantity of morpho-variations, which qualifies the species as one of the most diverse and complex taxa of the genus Pedicularis. In this study, the phylogeny of P. sect. Cyathophora was reconstructed based on four chloroplast genes (matK, rbcL, trnH-psbA, and trnL-F) and one nuclear gene, ITS, using 78 samples, with an emphasis on the P. rex complex. Also morphometric analyses were used to assess the morphological variation in P. rex complex and taxonomic revisions were presented for some taxa of this complex. Thirteen microsatellite markers were developed from three microsatellite-enriched libraries (AG, AC and AAG) of P. rex with a modified biotin-streptavidin capture technique, which could be used for further studies on the genetic diversity and population structure of this species and its congeners. Main results were summarized as follows: 1. Molecular phylogeny of P. sect. Cyathophora H. L. Li， The phylogeny of P. sect. Cyathophora was reconstructed based on chloroplast matK, rbcL, trnH-psbA, trnL-F and one nuclear gene, ITS, involving six species and 11 outgroups with a total of 78 samples. Our study showed that the monophyly of this group was strongly supported, in which P. superba was monophyletic based on chloroplast genes. The samples of P. cyathophylloides from Sichuan nested within P. cyathophylla. Ancient hybridization may occur between the two species. P. xiangchengensis is better to treat as a synonym of P. cyathophylla. Especially, P. ser. Reges was complicated, in which the individuals from different populations in the same species were rarely monophyletic. In addition, individuals forming monophyletic groups were not geographically close. In P. ser. Reges, lineage sorting of chloroplast DNA variations following rapid divergence is likely to have caused complex phylogeny of the taxa, which was reconstructed in our study. Hybridzation, introgression, lineage sorting and adaptive radiation may play important roles in the evolution of these taxa studied. Multiple, independent data sets are needed for resolving phylogenetic relationships of rapidly diverged lineages in P. sect. Cyathophora. 2. The taxonomic revision of P. rex complex，A total of 165 specimens with about 14 vegetative and eight reproductive characters were involved in the morphometric analysis. The results of PCA did not support six taxa for it emerged considerable morphological overlap. P. rex var. rockii was promoted to subspecies level for its prominent performance in PCA. Three taxa, P. rex subsp. pseudocyathus, P. rex subsp. zayuensis and P. rex subsp. parva, were merged into P. rex subsp. rex. Then P. rex subsp. lipskyana kept still at its rank for its purple red corolla color. The result of DA led to the selection of diagnostic traits between the P. rex and P. thamnophila. P. rex has less dissected leaves than P. thamnophila. P. rex is various in leaves number in whole, but P. thamnophila has stable three leaves in whole. Itpossesses a smaller corolla than P. rex. But P. thamnophila subsp. cupuliformis is a little different from P. thamnophila subsp. thamnophila in some vegetative characters, and it has purple stripes with the lower lip, so it is kept at the subspecies level. 3. Isolation and characterization of microsatellite loci from P. rex， We developed 13 microsatellite markers from three microsatellite-enriched libraries (AG, AC and AAG) of P. rex with a modified biotin-streptavidin capture technique. Polymorphism of each locus was assessed in 22 individuals with representation of five populations of P. rex. Additionally, among the 13 identified microsatellite markers, eleven of them were successfully amplified in species P. thamnophila, and five of them showed polymorphisms. This study may provide important information for further investigation on the population genetics, introduction and acclimatization of P. rex and its 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in the genus of Pedicularis L. have high values in horticulture and some species are medicinal plants. However, they have a reputation for being uncultivable, which is an obstacle for their exploitation and utilization. In this dissertation, the hemiparasitism of Pedicularis species was studied systematically for the first time, and the successful cultivation of three Pedicularis species (Pedicularis densispica，Pedicularis cephalantha and Pedicularis rex) throughout all life stages was achieved for the first time. With several representative species as main study materials, a series of experiments on seed germination, vegetation survey in Pedicularis communities, and pot cultivation were carried out. We studied Pedicularis parasitic habit systematically and aimed to achieve progress in the cultivation of Pedicularis species based on the understanding of their parasitic habit. The main results are as follows:1. The dormancy showed by the tested Pedicularis species was non-deep physiological dormancy. Seed dormancy could be overcome by moist-chilling and GA3. Cold stratification for 30-60 days, or 500-1000 mg/L GA3 were the optimal treatments for germination percentage, and stratification for 15 days, or 1000 mg/L GA3 were the optimal treatments for mean germination time. The combination of cold stratification and GA3 were more effective measures to promote seed germination. The optimal germination conditions varied with species, while as a whole, the highest germination percentages were obtained from treatments of 500-1000 mg/L GA3 followed by 15-30 days stratification, and the lowest values of mean germination time were obtained from treatments of 100-1000mg/L GA3 followed by 15-day stratification. Seed germination was not the obstacle for the cultivation of Pedicularis species.2. Host range and host selectivity of a Pedicularis species were studied systematically for the first time. Examinations of haustorial connections revealed that P. densispica had a wide host range, and it can form haustorial connections on the roots of 33 species belonging to 14 families. Compositae (8 species), Gramineae (5 species) and Leguminosae (5 species) species comprised major hosts. In addition, self-parasitism was observed. Haustoria were non-randomly distributed among host species, suggesting that there was some host selectivity. P. densispica generally preferred species in the families of Gramineae and Cyperaceae. The results of association analysis and correlation analysis based on vegetation survey supported the result of examinations of haustorial connections. And correlation analysis was a better way to test host selectivity.3. This is the first report for the performance of Pedicularis species in cultivation throughout all life stages (from seeds to seeds). The high dependence of Pedicularis specieson host plants and their host preference were demonstrated in this study. Pedicularis speciesstrongly depended on host presence, while host plants were essential to Pedicularis speciesnot for survival but for proper development. Different Pedicularis species preferred to different hosts. Host defoliation was a useful promoting measure for the cultivation of Pedicularis species. Pedicularis species reduced the performance of host plants. With the assistance of hosts, three Pedicularis species were cultivated successfully and they retained high horticulture 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a rare genus endemic to China, has two species S. grevilleoides and S. albofusca. To explore systematic position and endangerment mechanisms of Sinopteris, its sporophyte morphological characteristics, gametophyte development, cytology, molecular phologenetics, and biogeography were investigated based on the extensive investigation of its distribution, existing circumstances and ecological adaptation. The main progresses are briefly summarized as follows:1. Resource investigation, S. grevilleoides has a scattered distribution only in very few areas of Dayao, Binchuan, Qiaojia in Yunnan and Qingchuan in Sichuan; the elevation scope of its habitate is 1100-1800m. S. albofusca distributes in areas, 1700-2800 meters above sealevel in Hebei, Beijing, Yunnan, Sichuan, Guangxi, Hunan and Guizhou. Our investigation showed that the resources of the two species were decreasing and it was very difficult to find them in wild, even in distribution areas on record. 2. Sporophyte morphological characteristics, The plants of Sinopteris are evergreen small xerophytes. Fronds are texture coriaceous, pentagon, upper surface smooth, under surface whitish-farinose; veins pinnate branching, not seen above, but very prominently raised beneath. Sori are marginal, terminal, consisting of one or two large, globose, subsessile sporangium, provided with a very broad annulus. Indusia deeply cut into triangular, toothed lobes. The form of frond epidermis of S. grevilleoides and S. albofusca bring into correspondence with each other: the epidermis cells are irregular narrow strip with sinuous anticlinal walls; the stomatal apparatus exists on the lower epidermis and its types according to Dilcher’s [26] nomination are polycytic and axillocytic. Scales attaching to the base of petiole are brown, lanceolate.3. Gametophyte development, The spores of S. grevilleoides and S. albofusca were cultured in improved Knop’s agar medium and three kinds of soil substrata respectively. Spore germination and gametophyte development were observed. The impact of four culture substrata on the gametophyte development and sexual reproduction of S. grevilleoides was compared. The results are as follows:(1) The mature spores of Sinopteris were black-brown, isospory, regular tetrahedron, trilete, blunt triangle in polar view, scoop in equatorial view. Spore germination was of Vittaria-type and gametophyte exhibited Adiantum-type development. The adult prothalli were symmetric cordate. Antheridium protruding from the surface of prothalus was nearly spherosome. Archegonium was born near the notch on the ventral face of the cordate prothalus and those near the notch matured late. Mature archegonium was tall and slender, whose neck was composed of four lines cells and three to five layerscells each line. The four cells at the top separated when the archegonium matured. (2) The gametophyte and infant sporophyte morphological development of S. grevilleoides represented very different features on different culture substrata. Mature cordate prothalli on humus soil did not bear archegonias, and thus the process of sexual reproduction ofS. grevilleoides could not be finished. The substratum composed of humus soil and original soil in ratio of 1:1 was the most suitable substratum for gametophyte development and sexual reproduction of this species. Infant sporophytes needed extra nutrient solution to sustain its growth on improved Knop''s agar medium.(3) The main contaminants were bacteria, fungi and algae during sterile cultures and algae, moss and other higher plants during soil cultures.4. Cytology, Chromosome numbers of S. grevilleoides and S. albofusca were investigated firstly. The result showed that the chromosome numbers of the two species both were 2n=60 and the basic choromosome number was x=30. 5. Molecular phologenetics, The phylogeny of 26 species was estimated from combined analyses of four cpDNA sequence data sets (rps4, rps4-trnS, atpB, atpB-rbcL and trnL-F). Most parsimonious (MP) and Bayesian analysis both shows the genus Sinopteris is monophyletic with strong support and it is nearly related to A. subargentea from Ser. Argentea of Aleuritopteris. The MPsystematic tree also shows: both Leptolepidium and Cheilosoria are not monophyletic; Cheilosoria is nearly related to Ser. Argentea of Aleuritopteris and Leptolepidium is nearly related to Ser. Farinosae of Aleuritopteris.6. Population and community structure, The two species of Sinopteris mainly grow in the crevices of cliffs and the numbers of individuals within populations are small. In the community, they are less competitive and under dog. Its geographical distribution requires strict elevation, humidity and illumination. S. grevilleoides often grows in the brushwood or grass. Thickets of S. albofusca usually grows in the brushwood or coniferous forest. 7. Soil property,S. grevilleoides grows in purplish soil in wild; the soil is very thin and it isdifficult to store water. S. albofusca grows in sticky and infertile red soil. Analysis of the soil chemical property shows: the two original soils both are acid soil with high calcium and available B; their total Pand K is low.8. Endangerment factors and conservation measures, Endangerment factors: (1) Their distribution areas are so narrow that they are difficult to cope with environmental changes. (2)Their own physiological and ecological characteristics: spore germination was exigent over humidity, illumination and soil; their ecological circumstances cause their rhizomes to have no too much room to extend. Based on the previous two, reproduction by spores and rhizomes for the genus Sinopteris is limited. (3)To some extent, human-induced habitat loss, accompanying habit fragmentation, and natural disaster such as drought and fire accelerated the endangerment process. Conservation measures: (1) Have a clear and definite acquaintance to recent population size, distribution and growth dynamics by strengthening field investigation. (2) According to the specific circumstances, introduce in-situ and ex-situ conservation. (3)Rejuvenate and expand Sinopteris by adopting division propagation and artificial reproduction by spores. (4) Exploit species of the genus as greening and floral 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