955 resultados para floral odour
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At first it was intended to confine the collections to the Canal Zone proper, but as the faunal and floral areas extended to the north and south of this region, it was decided to carry the work into the Republic of Panama, a step which met with the hearty approval of that Republic. The work accomplished has been very valuable to science, including collections and observations of vertebrate animals, land and fresh water mollusks, and plants, including flowering plants, grasses and ferns. Special attention will be given during the coming season to vertebrate animals, insects, crustaceans, rotifers and other minute freshwater animals, and also to the microscopic plants known as diatoms. Includes appendix of papers that resulted.
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Raw fillets of saithe, red fish and salmon were stored over a period of 2 days at 6 to 9 °C before preparing by frying, deep-frying or cooking in a microwave oven. The raw and prepared fillets were tested for bacterial loads, TVBN (total volatile basic nitrogene) and for the sensorial status. It was shown that saithe and red fish started more rapidly to spoil under these conditions than salmon. The fillets showed growing bacterial populations and produced high amounts of TVBN. Sensory changes, especially in flavour and odour, took place but the scores kept in an acceptable range. The investigations indicated that it is possible to store raw fillets for 1 or even 2 days in refrigerator but due to the loss of quality it is recommended to use only really fresh fillets stored in very clean containers. It was further shown that it is very important to heat fillets sufficiently during cooking or frying in order to destroy all microorganisms and to obtain safe and stable meals.
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Baltic Sea herring is a traditional raw material for the German fish processing industry and the fresh fish market. This applies also for the spring herring of the spawning population of the waters around the island of Rügen. Reduction of the fat content to about 5 % during the spawning cycle limits the processing possibilities of mature herring from this area. Failures in taste and odour (tainting), a common problem of the past have not been detected in the last 3 years. Infestation by nematodes are comparable to other herring stocks and contamination levels of organic and inorganic contaminants are well below allowable limits. The annual German fishing quota of about 85000 t of Baltic Sea herring is now utilised only to 10 %. For a stronger utilization of this stock as in the 70th and 80th , there are scarcely prerequisites. The project of a central processing plant on the island Rügen for about 50000 t of herring as raw material is not realistic. The answer to the question asked at the beginning of this article, whether Baltic Sea herring represents a raw material for the German fish processing industry, is YES, dispite some restrictions.
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The phytoplankton distribution of the Shen Reservoir, Bukuru in the Jos Plateau, Nigeria was monitored at 6 depths. Higher floral abundance occurred within the upper 00-03 meters with highest values at the first 1 meter. Bacillariophyceae and Dinophyceae recorded higher values in March-April with lower values in July and January respectively. Phytoplankton were most abundant in the rainy season. Secchi disc transparency was lowest in the peak of the rainy season (July) due to higher levels of suspended matter resulting from the increased run-off from surrounding farmlands of allochthonous materials as well as higher levels of phytoplankton population arising from the former factor. The low water temperature of December/January 15 degree C plus or minus 2 degree C might have depressed growth among the major groups of plankters but enhanced rapid multiplication of the Chlorophyta, Trachelomonas which showed a bloom at this season
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This study was carried out to assess consumers' acceptance of kilishi prepared from Labeo coubie and Hyperopisus bebe occidentalis in Sokoto. The organoleptic properties (texture, odour, taste and flavour) of kilishi in its fresh form and under storage for 16 weeks were determined. The mean scores for the organoleptic assessment (6.90 and 7.19 for kilishi of Labeo and Hyperopisus respectively) showed that fish kilishi was highly acceptable. Hyperopisus kilishi recorded slightly higher mean scores for the tested organoleptic properties. The declining pattern of the sensory assessment scores with length of storage indicated that the optimum storage period under the room temperature for kilishi made from the experimental fish species in the study area was 6-8 weeks. Further research on appropriate storage methods is desirable. However, preparation of fish kilishi could be explored as alternative preservation technique to reduce fish spoilage especially during the glut in supply and to diversify fish products
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During the 160th research cruise of the FRV "Walther Herwig III" in the North Sea in May 1995 an ice-storage experiment with whiting was performed. Gutted whiting with and without spleen was stored in melting water-ice. Freshness and/or spoilage were monitored by measuring sensory, chemical, physical and microbiological indicators. It was found that besides the classical sensory assessment on the cooked sample and the EU-quality grading scheme, the microbiological counts were of major importance for the determination of the degree of freshness or spoilage. The cfu (colony forming units) of spoilage bacteria on the skin correlated significantly with time in ice. A very good correlation was also found for the cfu of spoilage bacteria with the sensory assessed odour of the cooked fillet sample. The measurement of the fish tissue with the Intellectron Fischtester VI and the determination of the creatine content in fillet are both suitable freshness and spoilage indicators. The pH-value measured in different body compartments and in musele homogenate and the ammonia content are only of limited value for freshness determination. Removal of kidney did not influence the shelf life.
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Tastes and odours are amongst the few water quality standards immediately apparent to a consumer and, as a result, account for most consumer complaints about water quality. Although taste and odour problems can arise from a great many sources, from an operational point of view they are either ”predictable” or ”unpredictable”. The former - which include problems related to actinomycete and algal growth - have a tendency to occur in certain types of water under certain combinations of conditions, whereas the latter - typically chemical spills - can occur anywhere. Long-term control is one option for predictable problems, although biomanipulation on a large scale has had utile success. Detection and avoidance is a more practicable option for both predictable and unpredictable problems, particularly if the distribution network can be serviced from other sources. Where these are not feasible, then water treatment, typically using activated carbon, is possible. In general there is a reasonable understanding of what compounds cause taste and odour problems, and how to treat these. An efficient taste and odour control programme therefore relies ultimately on good management of existing resources. However, a number of problems lie outside the remit of water supply companies and will require more fundamental regulation of activities in the catchment.
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A vegetação da Ilha Grande faz parte do Bioma Floresta Atlântica, que possui altos índices de biodiversidade e cobre amplas regiões de zonas climáticas e formações vegetacionais tropicais a subtropicais. No Brasil, estende-se numa estreita faixa ao longo de quase toda a costa atlântica e interioriza-se atingindo parte da Argentina e do Paraguai. Asteraceae é a terceira maior família em número de espécies na Floresta Atlântica. Assim, buscou-se conhecer a representatividade dessa família na Ilha Grande, objetivando contribuir com a política de preservação e manutenção de seus ecossistemas. Nesse contexto, promoveu-se um levantamento bibliográfico, consultas a herbários e excursões periódicas de coleta em campo. O material coletado foi depositado no herbário da Universidade do Estado do Rio de Janeiro (HRJ). Registrou-se na área de estudo 67 espécies subordinadas a 37 gêneros. Os gêneros são a seguir denominados: Achyrocline (3 spp.), Adenostemma (1sp.), Ageratum (1 sp.), Austrocritonia (1 sp.), Astroeupatorium (1 sp.), Baccharis (8 spp.), Bidens (1 sp.), Blainvillea (1 sp.), Centratherum (1 sp.), Chaptalia (1 sp.), Chromolaena (3 spp.), Conyza (1 sp.), Cosmos (1 sp.), Eclipta (1 sp.), Elephantopus (2 spp.), Emilia (1 sp.), Erechtites (1 sp.), Galinsoga (1 sp.), Gamochaeta (1 sp.), Grazielia (1 sp.), Heterocondylus (2 spp.), Mikania (13 spp.), Piptocarpha (2 spp.), Pluchea (1 sp.), Praxelis (1 sp.), Pseudogynoxys (1 sp.), Pterocaulon (1 sp.), Sphagneticola (1 sp.), Sonchus (1 sp.), Steymarkina (1 sp.), Struchium (1 sp.), Synedrella (1 sp.), Tilesia (1 sp.), Tithonia (1 sp.), Trixis (1 sp.), Verbesina (1 sp.) e Vernonia (5 spp.). Estes gêneros estão abrigados sob nove tribos. São citadas pela primeira vez para o Estado do Rio de Janeiro as espécies Mikania campanulata e Struchium sparganophorum.
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A subtribo Elephantopinae tem sido alvo de poucos estudos taxonômicos e palinológicos, necessitando de uma reavaliação em seus limites. Assim, o presente estudo teve como proposta atualizar e ampliar o conhecimento de 13 espécies de Elephantopinae ocorrentes no Brasil através de um acurado estudo palinológico, da anatomia foliar e taxonômico, promovendo a delimitação das espécies dos gêneros Elephantopus (10 espécies) Orthopappus (uma espécie) e Pseudelephantopus (duas espécies)além de oferecer subsídios para posteriores análises filogenéticas. O material botânico utilizado foi obtido através de exsicatas depositadas nos herbários brasileiros e de material coletado em campo. Os grãos de pólen foram tratados pelo método acetolítico, sendo posteriormente mensurados, descritos, e analisados sob microscopia de luz e eletrônica de varredura. As eletromicrografias foram obtidas de grãos de pólen não acetolisados, as análises taxonômicas baseiam-se na metodologia clássica. A anatomia foliar deu-se através da metodologia usual. Os resultados mostram grãos de pólen médios, 3-porados, de exina equinolofada com malhas poligonais organizadas ou não em Elephantopus e Pseudelephantopus, podendo apresentar interrupção na malha poral em Elephantopus. Em Orthopapus os grãos de pólen são 3-colporados, de sexina subequinolofada. A anatomia foliar possibilitou a separação de E. hirtiflorus e E. riparius através da forma ou contorno da nervura principal (planoconvexo), nas demais espécies o contorno é biconvexo. As espécies apresentaram tricomas capitados bisseriados com 8 células, tricomas filamentosos 3-4 celulares, bem como filamentosos de célula distal globosa ou ovóide em E. biflorus, E. micropappus, E. tomentous e Orthopappus angustifolius. Registrou-se variação na ornamentação da cutícula podendo ser lisa ou estriada entre as espécies, bem como a presença de substâncias pécticas na epiderme e drusas nos parênquimas. Em relação aos microcaracteres florais, foram considerados de importância diagnóstica os lóbulos da corola, que variaram entre glandular, glabro e penicilado, este ultimo apenas em E. hirtiflorus, os ápices das anteras que variaram entre obtuso, retuso em E. hirtiflorus e apiculado, lancelolado em E. mollis e E. racemosus, a base da antera variou entre sagitada, caudada lisa em E. riparius e E. tomentosus e caudada em E. riparius. Os macrocaracteres que apresentaram maior valor taxonômico foram a cipsela e o papus, a organização das coflorescências, o número de flores por capítulo e limbo foliar. Neste trabalho aceitou-se a segregação dos três gêneros, com base nos caracteres analisados
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Asteraceae é a segunda maior família de angiospermas com ca. 25.000 espécies distribuídas por todo mundo. Praxelis é o segundo maior gênero em número de espécies da subtribo Praxelinae, se caracteriza principalmente por apresentar capítulos cilíndricos/campanulados, receptáculo cônico e glabro, cipselas com carpopódio assimétrico e pápus com cerdas numerosas e barbeladas. No Brasil, as espécies predominam no Cerrado. O objetivo do presente trabalho foi analisar a palinologia e a taxonomia das espécies de Praxelis Cass. ocorrentes no Brasil. O material botânico utilizado foi obtido através de exsicatas depositadas nos herbários brasileiros. Os grãos de pólen foram acetolisados, medidos, descritos e ilustrados sob ML. Para observar detalhes da superfície e abertura, grãos de pólen não acetolisados foram analisados em MEV e, posteriormente, eletromicrografados. As peças florais foram colocadas sobre os mesmos suportes metálicos cobertos com fita dupla-face de carbono, previamente numerados. Para análise em MET, os grãos de pólen foram corados em cacodilato tamponado 0,125% OsO4, concentrados em pastilhas de ágar, e foram embebecidos em resina Epon-Araldite. As folhas, a capitulescência, o indumento do pedúnculo e o número de brácteas involucrais foram atributos relevantes para caracterizar as espécies, sobretudo o tamanho do invólucro e sua consistência. Foram caracterizados os grãos de pólen quanto à forma, ao tamanho, ao tipo de abertura, à polaridade e à constituição da exina. Foram descritos como: pequenos a médios, isopolares, oblato-esferoidais a subprolatos, 3(4)-colporados, com margem ornamentada, endoabertura com ou sem constricção, com presença de cávea, a sexina é espinhosa e sempre maior do que a nexina. Portanto, as análises palinológica e taxonômica foram utilizadas para segregar as espécies de Praxelis, porém apenas os atributos morfológicos externos foram mais informativos para diferenciar as espécies desse gênero.
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小檗科是毛茛类的核心成员之一,属于较原始的被子植物类群。目前有关该科在毛茛类的系统位置还没有一致的意见。一种观点认为它与毛茛科有密切关系,属于本目中较原始的一支;另一些学者则根据小檗科与罂粟科在心皮结构上的相似性提出它们有较近的亲缘关系,并可能在毛茛类中处于较高的演化水平上。小檗科的范畴与科内系统至今仍存在着较大争论。自本世纪初以来已有若干系统发表,其中有的采取广义小檗科的概念,而有的则分别成立2-4个科。在生殖结构演化方面,有关小檗科植物花的一些形态学本质,如花基数、蜜腺来源、心皮性质等仍有待揭示。因此,有必要对该科的结构、分化和系统发育进行深入的研究。 本研究首次报导了小檗科9属植物的花部形态发生、10属植物胚胎发育、9属植物的种皮纹饰和八角莲属与桃儿七属的rbcL基因全序列。并综合已有的研究资料,对小檗科的系统学问题进行了讨论。主要实验结果及结论如下: 1花部形态发生 小檗科植物典型花部形态为三数轮列、雄蕊与花瓣对生、单生心皮、侧生胎座。花器官发育的独特性状包括雄蕊与花瓣以共同原基方式发生;心皮原基环形、周缘组织等速向上发育而建成瓶状心皮,不形成腹缝线。我们认为,三基数在小檗科植物中可能是共同起源的,为典型轮列花;花瓣(蜜腺叶)来源于花瓣一雄蕊共同原基,而小檗属、十大功劳属、兰山草属花瓣上的小囊状蜜腺是其本身的附属结构,与雄蕊无关;雄蕊与花瓣对生来源于雄蕊一花瓣共同原基的发生方式;小檗科的心皮发生方式在毛茛类中是独特的,它不同于Hell (1981,198 3)描述的囊状心皮. 2胚胎学 小檗科植物的主要胚胎学性状有:腺质绒毡层,药室内壁带状加厚,2细胞花粉粒,胚珠具双珠被、厚珠心,珠孔由内外珠被组成,直线型大孢子4分体,蓼型、稀待宵草型(红毛七属)或葱型(桃儿七属)胚囊,反足细胞宿存,核型胚乳,柳叶草型或茄型胚胎发生。根据小檗科各属植物的胚胎学与种皮表面雕纹性状的分化,小檗科可以划分为:南天竹属,小檗属一十大功劳属,红毛七属一牡丹草属一狮足草属,山荷叶属一八角莲属一桃儿七属一足叶草属,淫羊藿属一鲜黄连属5个属群。说明胚胎学性状对揭示本科植物属间及属群间系统关系有较大的价值。 3分子数据 利用本研究测得的八角莲属、桃儿七属rbcL基因序列,结合从GenBank中得到的小檗科其他9属、毛茛科(4属)、防己科(2属)和木通科(1属)的rbcL序列,用PAUP软件进行分支分析。支持广义小檗科是单系类群的观点;.山荷叶属、八角莲属和桃儿七属构成的分支得到Bootstrap分析的完全支持(100%),3属植物是科内的一个自然类群;具x=6染色体基数的草本属间存在密切关系(90%);十大功劳属与兰山草属近缘(73%);南天竹属与红毛七属可能有一定联系(53%)。 4系统排列 广义小檗科(17属)为一单系类群得到本研究所获得的花形态发生、胚胎学和分子证据支持;综合各学科的研究结果提出小檗科分5族系统的建议.其系统排列为:1).南天竹族,含南天竹属1属.2).小檗族,分小檗亚族(小檗属、十大功劳属)和兰山草亚族(草本的兰山草属),共3属.3).狮足草族,含红毛七属、狮足草属和牡丹草属3属。4).足叶草族,分足叶草亚族(足叶草属、八角莲属与桃儿七属)和山荷叶亚族(山荷叶属),共4属.5).淫羊藿族,分淫羊藿亚族(淫羊藿属、 Vancou ver ia、Jef fersonia、鲜黄连属与Bongardia)和裸花草亚族(裸花草属),共6属。 5系统位置 根据小檗科花形态发生的独特性状,如心皮瓶状发育、雄蕊一花瓣以共同原基发生、二者对生、花药瓣裂和分子系统学的结果,以及据已有资料比较,小檗科在毛茛目中的特征较独特,与毛茛科的关系较远。支持小檗科提升为目的处理。
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传统的低等金缕梅类包含7个科,即领春木科(Eupteleaceae)、昆栏树科(Trochodendraceae)、水青树科(Tetracentraceae)、连香树科(Cercidiphyllaceae)、折扇叶科(Myrothamnaceae)、悬铃木科(Platanaceae)和金缕梅科(Hamamelidaceae)。这些科之间的谱系关系以及金缕梅科内部的谱系关系是研究被子植物系统和进化的主要问题之一。本研究的目的在于通过对现有的和本研究中产生的分子和发育学资料的分析来对这些谱系问题进行探讨,希望使我们对这些类群的认识提高到了一个新的水平。 本研究测定了包括连香树科、领春木科、悬铃木科、水青树科和金缕梅科内所有亚科代表共计14个种的rbcL基因,matK基因和18S-26S核核糖体(nrDNA)内转录间隔区(ITS)的序列,并结合原有资料对所有低等金缕梅类(除非洲分布的折扇叶科)及其相近类群进行了分支分析。主要结论包括;传统低等金缕梅类是一个复系类群;金缕梅科也是一个复系类群;ITS序列分析支持将连香树科包含在金缕梅科内,但是,rbcL资料和matK资料不支持将连香树科包含在金缕梅科内。本研究支持金缕梅科内6个亚科的分类系统,即将其分为阿丁枫亚科(Altingioideae)、红花荷亚科(Rhodoleioideae)、马蹄荷亚科(Exbucklandioideae)、壳菜果亚科(Mytilarioideae)、双花木亚科(Disanthoideae)和金缕梅亚科(Hamamelidoideae),但壳菜果亚科的山桐柴属(Chunia)应置于马蹄荷亚科内。 本研究首次描述和报道了金缕梅科5个亚科6个种的花的形态发生。通过对本研究发现的新证据和文献中记载的资料分析表明金缕梅科的花发育的特征无明显的共同式样。从本研究提供的花发育资料得出的值得注意的结论是红花荷属在金缕梅科中的亚科地位得到支持。对牛鼻栓(Fortunearia sinensis Rehd. et Wils.)的花发育研究表明其花性分化在发育晚期才出现。 本文首次对显脉红荷(Rhodoleia henryi Tong)的胚胎学进行了研究。尽管红花荷属是金缕梅科内适应鸟传粉的特化类群,但其胚胎学的特征与金缕梅科其它植物并无明显区别。因此,本研究不支持红花荷属独立成科而支持其在金缕梅科内的亚科地位。
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千里光族是菊科中最大的一个族,约含有120属3400多种,全世界分布。千里光族的族下分类与系统发育一直是菊科研究中的难点。Jeffrey&Chen(1984)和千里光亚族(7属)。自此以后,有关东亚款冬亚族的范畴、亚族内各属的系统位置与亲缘关系就成了千里光族系统学研究中争论的焦点,至今仍未有定论。陈艺(1997)在东亚款冬亚族内发表假橐吾属新属。根据Jeffrey&Chen(1984)的概念,东亚款冬亚族现含13属。东亚款冬亚族目前存在的系统学问题主要有:(1)毛冠菊属是否与款冬亚族内的款冬属及蜂斗菜属近缘;(2)狗舌草亚族所辖3属是否是款冬亚族的自然成员;(3)东亚款冬亚族13属的系统位置及其亲缘关系;(4)东亚款冬亚族的影响范畴。 针对上述问题,本文以发现新的系统学证据为主要目的,较为全面地研究了东亚款冬亚族及其相关类群的外部形态学,首次报道了18属45种植物的花部微观性状,19属64种105个居群的核形态学特征,10属15种植物花粉壁的超薄结构及13属28种植物花粉的外壁纹饰,3种植物的胚胎发育和6种植物的ITS基因序列;并对所获得的证据进行了比较与综合分支分析。根据所获系统学资料,对东亚款冬亚族的上述系统学问题进行了讨论。主要实验结果及结论总结如下。 1. 外部形态 较为全面地观察与分析了东亚款冬亚族及其相关类群外部形态性状及其演化趋势,指出根状茎分枝方式、幼叶出土性状、花部茎基部叶是否宿存、叶柄是否纤维状宿存、花序托空心或实心、花药基部结构等不被以前作者重视或忽视的形态学性状可能具有重要的系统学价值。外部形态上,毛冠菊属与千里光族内的千里光属与款冬属等类群没有亲缘关系,而与紫菀族存在一定的渊源;狗舌草亚族则介于千里光亚族和款冬亚族之间;东亚款冬亚族13属可依据外部形态划分为6个属群。 2. 花部微观性状 东亚款冬亚族及其相关类群的花部微观性状主要包括:花药顶端不育组织、花药药室内壁、花药基部、花药领、花柱分枝顶端、花柱毛被、花柱分枝内表面构型、果柄细胞列数等。花部微观性状研究表明:毛冠菊属似乎应作为紫菀族中的一个孤立属;狗舌草亚族在某些衍征上与千里光亚族接近,而它的原始性状则类似于款冬亚族;根据花部微观性状对东亚款冬亚族的13个属划分出的自然属群与形态性状分析所得结果基本一致。 3. 核形态学 毛冠菊属的染色体基数为x=9,与紫菀族相同:而与千里光族和旋覆花族的染色体x=10存在显著的区别。狗舌草亚族中新发现2n=24、60、72三个染色体数目,亚族内染色体基数之间的演化关系不清楚,利用染色体基数来探讨狗舌草亚族所辖属的系统位置和属间关系仍还比较困难。东亚款冬亚族的染色体基数主要有x=30、29、28、27、26;款冬亚族中普遍存在的染色体基数x=30可能有3种来源;核型可分为2A、3A、2B三大类型,虽然三种类型之间的染色体基数(x=30)有可能是不同起源的,但从整个千里光族来看,3A、2B应是较进化的类型。根据核形态学特征,东亚款冬亚族可分为款冬型、蟹甲草型和大吴风草型三类。在蟹甲草型染色体进化支上,存在染色体的非整倍性下降趋势。东亚款冬亚族存在的种内多倍性可能与不同的生境有关。 4. 花粉学 通过对东亚款冬亚族及其相关类群的花粉壁超微结构与扫描电镜观察,发现花粉壁超微结构存在“千里光型”和“向日葵型”两种类型。扫描电镜下,花粉外壁均为刺状纹饰,但在刺的长短、刺部是否膨大、刺基膨大上的纹饰以及刺基之间的纹饰等方面存在差别。毛冠菊属的花粉性状组合支持将其置于紫菀族中。狗舌草亚族的花粉性状与款冬亚族的有些属相似。款冬亚族各属的花粉生状组合所划分的属群与形态学、核形态学等方面的研究结果较为一致。 5. 胚胎学 款冬、掌裂橐吾和珠毛蟹甲草共同拥有药壁发育双子叶型;绒毡层发育属“The Comos bipinnatus”型;成熟花粉为3细胞型;单珠被,薄珠心,倒生胚珠;具发育相似的珠被绒毡层:胚橐发育4孢子型等胚胎学特征,但它们在胚囊发育方式、反足细胞数目、发殖物候上还存在区别。三属代表植物的胚胎学研究支持将它们作为同一亚族的成员。 6. 分子数据 利用本研究测得的华千里光属4种、狗舌草属1种和大吴风草属的ITS-1和ITS-2基因序列,结合从Genbank中得出的千里光族5属8种植物得ITS序列,用PAUP软件进行分析。研究结果表明,染色体基数、花粉超微结构、叶型和叶脉等性状在ITS分支图上存在趋同进化;华千里光属和狗舌草属不是单系群。并讨论了一些分类群的亲缘关系。 7. 分支分析 选择东亚款冬亚族和狗舌草亚族各属的代表种作为终端代表类群进行分支分析;分支分析表明染色体基数、与头状花序有关的性状、叶型及叶脉等性状存在较多的平行与逆转进化,而根状茎分枝类型、花部微观性状以及花粉性状则表现出较大的一致性。根据分支结果,作者认为应建立包括狗舌草亚族在内的广义东亚款冬亚族。 8. 系统学 毛冠菊属的性状组合支持将其置于紫菀族,排除在千里光族之外。广义东亚款冬亚族可分为7个属群:蟹甲草属群(包括蟹甲草属,华蟹甲草属,小蟹甲草属,兔儿伞属,蟹甲木属)、毛柱菊属群(毛柱菊属)、橐吾属群(包括橐吾属,垂头菊属,假橐吾属)、狗舌草属群(狗舌草属,华千里光属,羽叶千里光属)、多榔菊属群(多榔菊属)、大吴风草属群(大吴风草属)、款冬属群(款冬属,蜂斗菜属)。总结与讨论了各属群的系统位置、亲缘关系与存在的分类学问题。描述了华千里光属1新种。
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五味子属隶属于木兰亚纲八角目五味子科,全属共15种,呈东亚一北美间断分布,东亚是其分布中心。五味子科因花部器官数目多而不定,螺旋状排列等特征,被确认为原始多心皮类的一员,与八角科、木兰科、Winteraceae等有亲缘关系。同时,因为它具有一些特化的性状,如具3沟或异极6沟型花粉、花单性,而常被视为木兰亚纲中进化水平较高的类群。然而,近年来分子系统学的研究结果显示,五味子科是被子植物最基部的分支之一。这使得该科成为解决原始被子植物起源与演化问题的重要类群之一,而倍受关注。中国是五味子属植物分布最集中的地区,有着得天独厚的资源优势。在论述了该属研究现状和存在问题的基础上,作者重点从花发育形态学和分子系统学,对该类群进行了较为深入的研究,发现了一些重要的新的性状,为该类群的系统发育分析提供了有力的证据。 1. 借助于扫描电镜,对全属成熟雄花的6种变异类型的代表种,进行了全面细致的花形态建成的观察和分析。首次发现该属雄花形态建成的式样有3类,并分别将之命名为柱托型、平托型和球托型,它们的主要特征如下:柱托型在花发育的全过程里,花托部分保持柱状,雄蕊始终螺旋状排列;平托型的花原端在分化出雄蕊原基后,向上的生长减弱而侧向的生长增强,最终形成扁平五边形的花托,雄蕊先螺旋状排列后轮状排列:球托型的雄蕊原基仍是螺旋状向顶发生,但与此同时,雄花花托部分球形膨大,并逐渐在雄蕊着生处产生浅凹和孔穴。显然,这3种式样在花形态发生的早前期相一致,而在雄蕊原基萌动后并开始分化时,它们之间发生分异,导致成熟雄花的形态显著不同。作者认为,雄花形态式样反映了五味子属系统发育的渊源,柱托型是原始类型,平托型和球托型是衍生类型,它们可能在系统发育的较早期就已分异,各自朝不同的方向演化。根据这一新的证据,将五味子属分成3亚属:大花五味子亚属、五味子亚属和团蕊五味子亚属。它们雄花的形态发生式样分别为柱托型、平托型和球托型。还发现在雄花的形态发生同为柱托型的类群中,大花五味子群和少蕊五味子群的雄蕊在成熟时,药室由初始时位于药隔两侧的位置逐渐外转至药隔的背面或侧背面,造成开裂时呈外向或外侧向纵裂:而华中五味子群的雄蕊成熟后,药室仍位于药隔的两侧面,呈侧向纵裂。 南五味子属是五味子属最近缘的类群,以南五味子为代表种,进行了雄花形态发生的研究,该种雄花的形态建成式样属于柱托型,但又不同于五味子属的柱托型,其主要的区别表现在南五味子的雄蕊在成熟时变成轮状排列,该特征使它似乎界于大花五味子群和五味子群之间,呈过渡类型。因而,对于传统分类学中,依据果实的形态将五味子科划分为五味子属和南五味子属,需要重新审视。 2. 根据对花形态发生的观察和分析,文章提出五味子属中雄花形态发生式样为柱托型的大花五味子亚属是最原始的类群,而具平托型和球托型式样的类群演化程度都相对较高。五味子属雄花的演化趋势主要表现在花托从柱状向扁平状和球状演化:雄蕊数目由多数变为少数并最终向定数演化:雄蕊群由螺旋状排列向轮状排列演化。有性生殖过程中的传粉习性可能是导致五味子属雄花形态产生变异的原因。在白垩纪,鞘翅目昆虫仍是最主要的传粉媒介,它们在起到传粉作用的同时,也对植物的花器官造成严重损害。五味子属植物的祖先类群为了适应这种传粉习性,采取了不同的策略。雄花花托柱状,雄蕊螺旋状排列的类群,以多数雄蕊保证传粉得以完成:而在雄蕊数目少的类群中,则补偿性地增大花器官的营养组织部分,达到保护花药的目的。 关于五味子属植物单性花的性质,常被认为是较进化的性状。本研究结果显示,雄花中未出现雌性部分的残存,雌花中也未出现雄性部分的痕迹。因此,在形态上,五味子属表现出完全单性的特征。作者认为,五味子属的单性花性质不同于由两性花退化所形成的单性花,它可能反映了被子植物祖先类群花的另一种演化路线。 另外,作者通过对五味子属雌花形态发生的研究,比较和分析了诸多原始被子植物类群心皮的形态及其发生过程,赞成Endress认为离生多心皮类中既有对折心皮,也有囊状心皮,它们在原始类群中并存的观点。五味子属植物雌花的形态较一致,心皮在原基腹面基部的活动不明显,成熟心皮具明显的腹缝,腹缝两侧形成柱头脊,在类型上属于对折型心皮,体现了该属的原始性。 3. 作者选取了核基因组的rrs区和叶绿体基因组的trnL-F区,对五昧子属的10个种(代表了全部雄花类型)和南五味子属的4个种(分属2个组)以及作为外类群的华中八角,首次进行了序列分析。结果表明,从代表不同基因组的两个片段得到的分支图非常一致,都显示出:a) 五味子属的类群分别聚在3个不同的支上,并且这3个支完全对应于花形态发生的3种式样:b) 南五味子属的成员与五味子群聚在一起并得到Bootstrap分析很高的支持率,说明它们可能是单系的。该结果支持基于花形态发生研究所得到的结论。作者认为过去的分类中,将五味子属和南五味子属分割成两个属可能是不妥当的,值得深入研究。 4. 通过花发育形态学和分子系统学的研究,结合其它方面的资料,作者认为传统分类依据果实形态划分成的五味子属和南五味子属,可能都不是严格的单系类群。南五味子属的类群与五味子属中雄花花托为扁平五边形的类群显示出有很强的相关性。将两个属共同处理,分为3个亚属,似较适宜。由于尚需对南五味子属扩大取样研究,目前暂不对它进行处理。本文仅对传统意义上的五味子属的系统处理如下: 五味子属Schisandra Michx. 亚属1大花五味子亚属 Subgen.1 Pleiostema (Smith)Law 组1大花五味子组 Sect.1 PleiostemaSmith 组2华中五味子组 Sec.2 Sinoschisandra(Law)Liu 组3少蕊五味子组 Secc.3 Maximowiczia(Rupr.)Nakai 亚属2五味子亚属 Subgen.2 Schismxtm 亚属3团蕊五味子亚属 Subgen.3 Sphaerostema(Bl.)Law 5. 五味子科(或包括南五味子属在内的广义五味子属)现代分布格局显示,中国华中、华东和西南至印度支那北部的地域是该类群的多样性中心,而在整个分布区的外围,则零星散布着一些演化水平较高的类群。因此,五味子科很可能是在东亚大陆起源、分化后,往四周扩散。通过白令陆桥或欧美大陆到达北美:受第四纪冰川影响,形成东亚一北美间断分布。在东喜马拉雅、印度尼西亚的分布也都是扩散的结果。
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本文研究了茄科天仙子族7属以及有亲缘关系的颠茄属、茄参属和枸杞属的形态学、解剖学、孢粉学、种子形态学、胚胎学、花器官发生、分子系统学和分支系统学;论述了它们的结构、分化和系统关系。主要内容包括: 1.解剖学 在光学显微镜和扫描电镜下,观察了茄科天仙子族(Hyoscyameae)7属21种及邻近4属4种共25种植物的叶表皮特征,结果表明:气孔器在各种植物的上、下的表皮均有分布,多为无规则型,也有不等细胞型;叶表皮细胞形状有不规则形、十字形、近长圆形和不规则多边形,垂周壁较平直或各种波状。叶表皮气孔器外拱盖内缘近平滑、浅波状或波状;角质膜多为具条纹,也有同时具条纹和颗粒或鳞片,偶尔仅具颗粒和鳞片或光滑。气孔器类型可以作为区分部分属的依据之一。颠茄叶表皮细胞形状和垂周壁式样在一些属和种中有一定的规律性,并与外部形态、孢粉学性状或地理分布相关;叶片表面角质膜的特征对属和种的划分也有一定意义。 2.种子形态学 在扫描电子显微镜下首次全面研究了天仙子族及颠茄属和茄参属等共10属25种植物的种子形态学特征,结果表明: 该族植物种子表面纹饰有脑纹状或近脑纹状和网状两大类,除了山莨菪属Anisodus和Hyoscyamus.pusillus 外,其余类群均为网状。外种皮细胞形状和周壁式样有两大类,一类为多边形至近圆形,等径,周壁近平直,另一类为形状不规则,常不等径,周壁为各式波状;外种皮细胞周壁内侧有皱波状、瘤状、小颗粒、片状或小刺等各式附属物。各属植物的种子大小、形状及种脐着生位置的系统学意义不大。但是,种脐突出与否、种子外种皮细胞形状和周壁式样,尤其是周壁内侧附属物等性状具有较大的分类学意义。 3.孢粉学 在扫描电镜下研究了该族7属及其它3属共10属23种植物的花粉形态学特征。结果表明:天仙子族的花粉粒形状为球形、近球形、扁球形和长球形;萌发孔类型有无萌发孔、不规则的拟孔、3沟、4 沟、3- 4沟、散沟和3孔沟;外壁纹饰为小刺状、瘤状、各式条纹状、条纹-穴状、条纹-网状、皱波状、细网状和网状。沟膜近光滑、具小颗粒、具小刺状突起或具瘤状突起。各属植物的花粉形态在萌发孔的有无、萌发孔的类型和外壁纹饰等方面有较大的差异,可以作为探讨属间分类和系统关系的重要依据。天仙子族植物花粉萌发孔的演化趋势为:无萌发孔→ 3、4沟→3孔沟。无萌发孔的山莨菪属是原始类群,三孔沟的马尿泡和天仙子等属是进化类群。 4.胚胎学 本论文首次研究了天仙子族山莨菪Anisodus tanguticus的胚胎学特征,结果表明:山莨菪的小孢子形成为同时型,腺质绒毡层;胚珠半倒生,单珠被,薄珠心;蓼型胚囊;其胚胎发生类型可能为茄型;胚乳形成为细胞型。山莨菪的胚胎学性状与马尿泡十分相似。胚胎学证据支持山莨菪属与马尿泡属关系密切的观点。 5.花器官发生 首次全面研究了天仙子族的山莨菪属(山莨菪)、马尿泡属(马尿泡)、天仙子属(天仙子)及其邻近2属共5种植物的花器官发生特征,结果表明:山莨菪和天仙子的5个花萼裂片原基突起最早发生,且几乎是近同时环状发生,其排列方式从花发育早期直至花成熟后始终为轮状排列。而马尿泡和颠茄首先是5个花萼裂片原基突起依次螺旋状发生,但在其5个花萼裂片原基都出现后,其排列方式仍为轮状排列。不同之处是:马尿泡的花萼裂片原基在其花冠裂片原基刚刚不明显地发生时,就已经在基部联合并形成了一个环,而颠茄的花萼裂片原基直到雄蕊原基突起开始发生并较明显时,花萼裂片原基仍然是分离的,当花顶的中央开始出现很不明显的雌蕊原基的凹陷时,花萼裂片原基才开始联合。青海茄参与其它4种植物的不同之处在于柱头2裂或3裂,而后4者的柱头均为2裂。花器官发生的特征表明马尿泡与天仙子和山莨菪的关系较近。 6.基于叶绿体DNA trnL-F序列和核糖体DNA ITS序列的初步分析 以宁夏枸杞作为外类群,对代表天仙子族及其邻近3属的10种植物的ITS区和trnL-F序列进行了测定,将这两个序列结合构建系统树。将gaps作为fifth base处理时,马尿泡属和茄参属构成最基部的一支,赛莨菪属和天仙子属构成一支然后与泡囊草属构成姐妹群,另外一支由天蓬子属、颠茄属和山莨菪属组成,其中天蓬子属与颠茄属构成一支后与山莨菪属构成姐妹群。分子证据只是一个尝试,还不足以说明天仙子族各属的系统关系。 7.分支分析 以枸杞属为外类群,基于形态学、叶表皮、种子形态学、孢粉学等特征对天仙子族7属及颠茄属和茄参属进行了分支分析。结果表明:目前公认的天仙子族7属构成一个单系类群,而颠茄属和茄参属并没有包括在内,它们两者构成一支,并与天仙子族形成姐妹群的关系。这与该族传统的分类结果相一致。 8.天仙子族的分化和系统关系 基于上述研究结果,结合天仙子族各属的现代地理分布,讨论了各属的分化和系统关系。认为根据传统分类包括有7个属的天仙子族是一个自然的单系类群,目前还没有足够的证据说明是否应该将颠茄属和茄参属放在天仙子族中。