29 resultados para perianth


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Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

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In Arabidopsis, the identity of perianth and reproductive organs are specified by antagonistic action of two floral homeotic genes, APETALA2 (AP2) and AGAMOUS (AG). AP2 is also negatively regulated by an evolutionary conserved interaction with a microRNA, miR172, and has additional roles in general plant development. A kiwifruit gene with high levels of homology to AP2 and AP2-like genes from other plant species was identified. The transcript was abundant in the kiwifruit flower, particularly petal, suggesting a role in floral organ identity. Splice variants were identified, all containing both AP2 domains, including a variant that potentially produces a shorter transcript without the miRNA172 targeting site. Increased AP2 transcript accumulation was detected in the aberrant flowers of the mutant 'Pukekohe dwarf' with multiple perianth whorls and extended petaloid features. In contrast to normal kiwifruit flowers, the aberrant flowers failed to accumulate miR172 in the developing whorls, although accumulation was detected at the base of the flower. An additional role during dormancy in kiwifruit was proposed based on AP2 transcript accumulation in axillary buds before and after budbreak.

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花是被子植物特有的繁殖器官。花的发育取决于一个复杂的涉及到多个基因和过程的调控体系,因此花的起源和多样化过程实际上可以理解为这个调控体系的进化过程。所以,要全面地理解花和被子植物的起源和多样化,就必须研究花发育基因的功能和进化。 金粟兰科(Chloranthaceae)是基部被子植物的代表类群之一。与研究得比较深入的模式植物(如真双子叶植物中的拟南芥、金鱼草和矮牵牛等,和单子叶植物中的水稻和玉米等)相比,该科植物的花比较简单。花被以及雄蕊或雌蕊的丢失使得该科一些类群(如草珊瑚属Sarcandra和金粟兰属Chloranthus)具有被子植物中最简单的两性花(仅含一枚雄蕊和一枚雌蕊),而另一些类群(如Ascarina和雪香兰属Hedyosmum)具有了被子植物中最简单的单性花(雄花仅由一枚雄蕊、雌花由一枚雌蕊构成)。因此,对金粟兰科植物中花发育基因的研究不仅有助于理解花的起源和早期分化机制,还将为认识花部构造简单化的机制提供资料。 本研究以金粟兰(Chloranthus spicatus)为研究材料,从它的花和花序中分离得到了六个可能与花被的发生和发育有关的MADS-box基因,分析了它们的序列结构、系统发育关系、表达式样和进化中所受到的选择压力,探讨了金粟兰花发育和花被缺失的分子机理。主要研究结果包括: 1. 构建了金粟兰的花和花序的cDNA文库。构建工作使用了Clontech公司的SMART试剂盒,并采用其中的LD PCR方法,还使用了Stratagene公司的包装蛋白。该文库的初始滴度大约为5 × 106 pfu,重组率大约是90%, 插入片断几乎均大于0.5 kb。因此该文库质量优良,为以后的研究工作奠定了基础。 2. 从金粟兰的花中分离出了CsAP1、CsAP1a、CsAP1b、CsAP1c、CsAP3和CsSEP3基因。氨基酸序列分析结果表明它们都是MIKCc型MADS-box基因。系统发育分析结果表明CsAP1、CsAP1a、CsAP1b和CsAP1c与AP1/SQUA类基因聚在一起,而CsAP3和CsSEP3分别与AP3/DEF类和SEP1/2/3/4类基因聚在一起。CsAP1b和CsAP1c可能与CsAP1a互为复制本。但是二者在序列上有异常之处,因此可能只有CsAP1a具有功能。从序列上看CsAP1和CsSEP3能够正常行使功能。CsAP3的C末端出现了一个由鸟嘌呤到胸腺嘧啶的点突变,因此paleoAP3基序不完整,这可能影响了它的功能。 3. 用原位杂交的方法分析了CsAP1、CsAP3和CsSEP3的表达式样。CsAP1在穗状花序分生组织(包括苞片原基)、花原基、雄蕊和心皮原基、雄蕊裂片、花粉囊、胚珠、珠被和胚囊中表达。CsAP3在穗状花序分生组织中不表达,在花原基上发生雄蕊的位置开始表达,进而在雄蕊原基、雄蕊药隔裂片和花粉囊中表达,却不在心皮原基和心皮上表达。CsSEP3在穗状花序分生组织中也不表达,而在花原基、雄蕊原基、药隔裂片、花粉囊、心皮原基和胚珠中表达。CsAP1的表达模式反映了A功能基因决定花分生组织特性的原始作用;CsAP3的表达模式体现了B功能基因在雄性器官中的固有表达,反映了该类基因在两性器官分化中的原始作用;CsSEP3的表达模式反映了E功能基因提供成花背景(floral context)的作用。 4. 分析了已知的金粟兰的花发育相关基因受到的选择压力。同大多数近缘同源基因相比,CsAP1、CsAP3、CsPI、CsAG1受到负选择并且其强度没有明显差异;CsAG2和CsSEP3受到了更强的负选择;CsAP1a则受到减轻了的负选择。该结果表明除了CsAP1a之外,其它基因的功能可能没有改变。 5. 综上所述,在无花被的金粟兰中,仍然存在着与花被发育相关的基因,并且它们的功能没有改变,这充分反映了花发育ABC模型的保守性。金粟兰中花被的缺失可能与这些基因的下游基因有关,也可能与其它途径相关。CsAP1的复制以及CsAP3的末端突变可能是花被缺失之后的结果,而不是花被缺失的原因。

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花是被子植物区别于其它植物大类群的最重要的特征,其形态多种多样。花的发育取决于一个复杂的涉及到多个基因和过程的调控体系,因此花的起源和多样化过程实际上可以理解为这个调控体系的进化过程。在被子植物的不同物种中,花发育相关基因的组成并不相同,且经历了不同的进化历史,这意味着这些基因可能以不同的方式调控花的发育。相对于核心真双子叶植物相对稳定的花形态结构而言,基部被子植物的花具有丰富的多样性。因此,对基部被子植物花发育相关基因的研究对于我们理解被子植物花的进化非常重要。 金粟兰科(Chloranthaceae)是基部被子植物的代表类群之一。与研究得比较深入的模式植物相比,花被、雄蕊或雌蕊的缺失,使得该科植物的花比较简单。因此对该科植物中花发育基因的研究不仅有助于揭示花的起源以及花多样性分化的分子机制,还将为认识花部构造简单化的机制提供资料。本文以金粟兰(Chloranthus spicatus)为实验材料,取得了以下研究结果: 1.花发育相关基因的克隆 应用5’/3’RACE的方法,我们从金粟兰不同发育阶段的混合花芽中克隆到了与花发育相关的MADS-box基因:CsPI、CsAG1和CsAG2。 2. 两个A类MADS-box基因表达式样的对比分析 在营养分生组织向生殖分生组织的转变中,花原基的形成,以及随后雄蕊、心皮、花粉、胚珠和胚囊的发育中,CsAP1-1和 CsAP1-2基因均表达。唯一不同之处在于,在花发育成熟期,CsAP1-1在外珠被也有表达,而CsAP1-2在外珠被处没有表达,而只在内珠被处表达。这一结果反映了基因重复事件发生后,两个基因在功能上也有了一些分化。 3. B类基因功能的保守性和多样性 通过转基因实验和蛋白质相互作用研究对B类基因的功能和作用方式进行了研究,得到以下结果:1)金粟兰CsAP3基因的C末端的点突变所造成的paleoAP3基元的部分缺失对该基因的功能没有决定性的影响;2)金粟兰paleoAP3型基因CsAP3所编码蛋白的C末端以及paleoAP3基元,与拟南芥euAP3型基因AtAP3所编码蛋白的C末端以及euAP3基元没有功能上的不同;3)金粟兰paleoAP3型基因CsAP3与拟南芥euAP3型基因AtAP3 的主要功能存在一定差异,前者主要参与雄蕊形成,而后者既参与雄蕊的形成也参与花瓣的形成; 4)CsPI基因所编码的蛋白可以与AP3类蛋白相互作用进而影响花瓣的形成,因此该基因在功能上是保守的。 4. 金粟兰CsAG1基因的序列结构和功能分析 通过序列结构分析发现,CsAG1属于C类基因,具有保守的AG I基元和AG II基元。过量表达实验分析表明CsAG1的功能与A类基因的功能是相拮抗的。 5. 各类MADS-domain蛋白间的相互作用 在前面工作的基础上,我们首次对金粟兰中各类MADS-domain蛋白间的作用方式进行了研究。酵母双杂交结果表明:1)C末端的完整性对于MADS-domain蛋白二聚体的形成没有影响; 2)去掉M区的CsAP3蛋白与CsPI蛋白都能够形成异源二聚体,同时它们又可以各自形成同源二聚体; 3)E类蛋白既可以和A类或C类基因产物相互作用,也可以同AP3和PI型蛋白相互作用,充分体现了E类基因产物作用式样的保守性; 4)金粟兰中,FUL-like型基因所编码的蛋白CsAP1-1与CsSEP3和CsAG1也能形成异源二聚体,这与核心真双子叶植物的euFUL型蛋白在作用式样上是非常相似的。然而,金粟兰CsAP1-1蛋白不能形成同源二聚体。 综合以上结果发现,在无花被的金粟兰中,仍然存在着A、B、C/D、E类花发育相关的基因。这些基因的功能与核心真双子叶植物中同类基因的相比,有些是保守的,比如CsPI基因可以参与花被的形成;但也有一些是不同的,比如CsAP3基因主要参与雄蕊形成而非花被形成过程。由此可以看出被子植物花器官的发育是一个非常复杂的调控过程,不同植物中的调控机理及进化历程可能是不同的。

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诱导风信子(Hyacnthus orientalis L.)同一花被外植体上不同部位细胞分化花芽,从外源激素的作用和内探激素的变化探讨细胞脱分化启动的原因,研究了不同外源激素的组合下同一花被不同部位花芽分化的诱导频率:测定了花被上、中,下三部位切块离体培养前后的内源IAA和Z+ZR的含量;在此之前研究了GC-MS.MIM内标法测定微量植物材料内源IAA含量的可行性. GC-MS.MIM内标法是测定微量(0.5-1g)植物材料内源IAA含量的一种比较理想的方法,所需材料量一般为0.58.这一方法的材料前处理采用粗提液用C18Sep-pak柱初步分离纯化.HPLC进一步纯化,能获得纯度高的样品,且操作简便,省时省力. 风信子同一花被不同部位细胞均能分化花芽.当培养基中附加2.0mg/l Zeatin或2.0mg/16-BAP时,随着外探IAA浓度从0升高到10.0mg/l.捆胞分化花芽的部位从花被下部向上部移动。 离体培养前后同一花被上、中、下三部位内源IAA和Z+ZR含量测定结果表明.风信子同一花被内源IAA含量是上部最高,下部次之,中部最低,而内源Z+ZR含量从上向下依次增加;在附加不同外源激素的MS培养基上培养3天后,同一花被上,中、下三部位内源IAA和Z+ZR含量部有一定的变化。

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以比较形态解剖学方法,对苔纲(Hepaticae)植物雌性生殖苞的结构及其不同成分起源进行了研究。首次把苔纲雌性生殖苞(Perichaetium)分为蒴帽型(Calyptra type)、蒴萼型(Perianth type)、鞘萼型(Coelocaule-peri-anth type)、茎鞘型(Coelocaule type)、蒴囊型(Perigynium type)、总苞型(Involucre type)和托生总苞型(Carpoceph-alum involucre type)等雌苞类型,并指明其中蒴帽型雌苞(Perichaetium of calyptra type of Hepaticae)最简单,为演化原始类群,托生总苞型雌苞(Perichaetium of carpocephalum involucre type of Hepaticae)演化最高级。还首次提出了雌苞演化示意图和"苞图式"表示苔类的雌苞结构。

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A study of inflorescence and flower development in 12 species from four of the six subgenera of Gunnera (Gunneraceae) was carried out. In the species of subgenus Panke, initiation of floral apices along the partial inflorescences is acropetal but ends up in the late formation of a terminal flower, forming a cyme at maturity. The terminal flower is the largest and the most complete in terms of merosity and number of whorls and thus it is the most diagnostic in terms of species-level taxonomy. The lateral flowers undergo a basipetal gradient of organ reduction along the inflorescence, ranging from bisexual flowers (towards the distal region) to functionally (i.e. with staminodia) and structurally female flowers (towards the proximal region). Our results show that the terminal structure in Gunnera is a flower rather than a pseudanthium. The terminal flower is disymmetric, dimerous and bisexual, representing the common bauplan for Gunnera flowers. It has a differentiated perianth with two sepals and two alternate petals, the latter opposite the stamens and carpels. Comparisons with other members of the core eudicots with labile floral construction are addressed. We propose vegetative and floral putative synapomorphies for the sister-group relationship between Gunneraceae and Myrothamnaceae. (C) 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160, 262-283.

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Background and Aims: Molecular phylogenies have suggested a new circumscription for Fabales to include Leguminosae, Quillajaceae, Surianaceae and Polygalaceae. However, recent attempts to reconstruct the interfamilial relationships of the order have resulted in several alternative hypotheses, including a sister relationship between Quillajaceae and Surianaceae, the two species-poor families of Fabales. Here, floral morphology and ontogeny of these two families are investigated to explore evidence of a potential relationship between them. Floral traits are discussed with respect to early radiation in the order. Methods: Floral buds of representatives of Quillajaceae and Surianaceae were dissected and observed using light microscopy and scanning electron microscopy. Key Results Quillajaceae and Surianaceae possess some common traits, such as inflorescence morphology and perianth initiation, but development and organization of their reproductive whorls differ. In Quillaja, initiation of the diplostemonous androecium is unidirectional, overlapping with the petal primordia. In contrast, Suriana is obdiplostemonous, and floral organ initiation is simultaneous. Independent initiation of five carpels is common to both Quillaja and Suriana, but subsequent development differs; the antesepalous carpels of Quillaja become fused proximally and exhibit two rows of ovules, and in Suriana the gynoecium is apocarpous, gynobasic, with antepetalous biovulate carpels. Conclusions: Differences in the reproductive development and organization of Quillajaceae and Surianaceae cast doubt on their potential sister relationship. Instead, Quillaja resembles Leguminosae in some floral traits, a hypothesis not suggested by molecular-based phylogenies. Despite implicit associations of zygomorphy with species-rich clades and actinomorphy with species-poor families in Fabales, this correlation sometimes fails due to high variation in floral symmetry. Studies considering specific derived clades and reproductive biology could address more precise hypotheses of key innovation and differential diversification in the order.

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Keeled flowers are characteristic of many species of Polygalaceae (tribe Polygaleae), in which there is often a distinctive crest located at the abaxial petal. This study of floral ontogeny across the entire family highlights potential suites of characters that relate to the evolution of keeled and crested flowers. One character suite encompasses interconnected transformations of the lateral perianth organs acting as an evolutionary module: bracteoles, lateral sepals (with delayed initiation and petaloid appearance), and lateral petals (suppressed or lost). The plastochron between initiation of the lateral sepals and that of the other sepals is relatively long in the tribe Polygaleae, in which the calyx is usually heteromorphic. By contrast, in the petal whorl, the difference between a zygomorphic and an actinomorphic corolla involves organ suppression rather than heterochrony. Four primary androecial patterns are identified in the family, and the gynoecium ranges between syncarpous‐bicarpellate and multicarpellate. Stigma diversity is based on two primary morphological types: one in which the papillate stigmatic surfaces lie close together, the other in which they are physically separated. The floral ontogeny of Polygalaceae is considered alongside comparative data available for other members of the order Fabales, and hypotheses to account for the similarities and differences between keeled flowers are discussed.

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• Background and Aims: Eriocaulaceae (Poales) is currently divided in two subfamilies: Eriocauloideae, which comprises two genera and Paepalanthoideae, with nine genera. The floral anatomy of Actinocephalus polyanthus, Leiothrix fluitans, Paepalanthus chlorocephalus, P. flaccidus and Rondonanthus roraimae was studied here. The flowers of these species of Paepalanthoideae are unisexual, and form capitulum-type inflorescences. Staminate and pistillate flowers are randomly distributed in the capitulum and develop centripetally. This work aims to establish a floral nomenclature for the Eriocaulaceae to provide more information about the taxonomy and phylogeny of the family. • Methods: Light microscopy, scanning electron microscopy and chemical tests were used to investigate the floral structures. • Key Results: Staminate and pistillate flowers are trimerous (except in P. flaccidus, which presents dimerous flowers), and the perianth of all species is differentiated into sepals and petals. Staminate flowers present an androecium with scale-like staminodes (not in R. roraimae) and fertile stamens, and nectariferous pistillodes. Pistillate flowers present scale-like staminodes (except for R. roraimae, which presents elongated and vascularized staminodes), and a gynoecium with a hollow style, ramified in stigmatic and nectariferous portions. • Conclusions: The scale-like staminodes present in the species of Paepalanthoideae indicate a probable reduction of the outer whorl of stamens present in species of Eriocauloideae. Among the Paepalanthoideae genera, Rondonanthus, which is probably basal, shows vascularized staminodes in their pistillate flowers. The occurrence of nectariferous pistillodes in staminate flowers and that of nectariferous portions of the style in pistillate flowers of Paepalanthoideae are emphasized as nectariferous structures in Eriocaulaceae. © The Author 2006. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved.

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Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

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Diese Studie befasst sich mit der Phylogenie und Biogeographie der australischen Camphorosmeae, die ein wichtiges Element der Flora arider Gebiete Australiens sind. Die molekularen Phylogenien wurden mit Hilfe Bayes’scher Statistik und „maximum likelihood”berechnet. Um das Alter der Gruppe und interner Linien abzuschätzen, wurden die Methoden „Nonparametric rate smoothing” und “penalized likelihood” benutzt. Morphologische Merkmale wurden nach Kriterien der Parsimonie auf den molekularen Baum aufgetragen. „Brooks parsimony analysis”, „cladistic analysis of distributions and endemism”, „dispersal-vicariance analysis”,„ancestral area analysis” und „weighted ancestral area analysis” wurden angewandt, um Abfolge und Richtungen der Ausbreitung der Gruppe in Australien zu analysieren.Von sieben getesteten Markern hatten nur die nukleären ETS und ITS genügend Variation für die phylogenetische Analyse der Camphorosmeae. Die plastidären Marker trnL-trnF spacer,trnP-psaJ spacer, rpS16 intron, rpL16 intron und trnS-trnG spacer zeigten kein ausreichendes phylogenetisches Signal. Die gefundenen phylogenetischen Hypothesen widersprechen der jetzigen Taxonomie der Gruppe. Neobassia, Threlkeldia, Osteocarpum und Enchylaena sollten den Gattungen Sclerolaena bzw. Maireana zugeordnet werden. Die kladistische Analyse der Fruchtanhängsel unterstützt die taxonomischen Ergebnisse der auf DNA basierenden Phylogenie. Allerdings hat die Behaarung, die bei anderen Gruppen der Chenopodiaceae als wichtiges taxonomisches Merkmal herangezogen wird, die Phylogenie nicht unterstützt. Vorfahren der heutigen Camphorosmeen sind im Miozän, vor ca. 8-14 Millionen Jahren, durch Fernausbreitung vermutlich aus Asien in Australien eingewandert. Anfängliche Diversifizierung fand während des späten Miozäns bis in das frühe Pliozän vor ca. 4-7 Millionen Jahren statt. Am Ende des Pliozäns existierten schon 45% - 72% der Abstammungslinien der jetzigen Camphorosmeen. Dies weist auf eine schnelle Ausbreitung hin. Das Alter stimmt mit dem Einsetzen der Aridisierung Australiens überein, und deutet darauf hin, dass die Ausbreitung der ariden Gebiete eine große Rolle bei der Diversifizierung der Gruppe spielte. Die Vorfahren der australischen Camphorosmeae scheinen die Südküste Australiens zuerst besiedeln zu haben. Dies geschah vor dem Einsetzen der Aridisierung des Kontinents. Die anschließende Ausbreitung erfolgte in verschiedene Richtungen und folgte der fortschreitenden Austrocknung im späten Tertiär und im ganzen Quartär. Durch ihre Anpassung an Trockenheit ist der Erfolg der Camphorosmeae in den ariden Gebieten zu erklären.Die Abwesenheit von klaren phylogenetischen und artspezifischen Signalen zwischen Arten der australischen Camphorosmeae ist auf das junge Alter und die schnelle Diversifizierung der Gruppe zurückzuführen, welche die Häufung von Mutationen und eine starke morphologische Differenzierung nicht zugelassen haben.

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The biostratigraphic classification of the Pleistocene in north-western and central Europe is still insufficiently known, in spite of numerous geological and vegetation-history investigations. The question is not even clear, for example, how often a warm-period vegetation with thermophilous trees such as Quercus, Ulmus, Tilia, Carpinus etc could develop here. In past years, on the basis of several geological and vegetation-history findings, suspicion has often been expressed that some of the classical stages of the Pleistocene could include more warm periods than heretofore assumed, and as a result of recent investigations the period between the Waal and Holstein interglacials seems to include at least two warm periods, of which the Cromer is one. This paper contributes to this problem. The interglacial sediments coming from the Elm-Mountains near Brunswick and from the Osterholz near Elze - both within the limits of the German Mittelgebirge - were investigated by pollen analysis. In both cases a Pinus-Betula zone and a QM zone were found. The vegetation development of the Pinus-Betula zone is characterized in both sequences by the early appearance of Picea. Because of strong local influence at the Osterholz a detailed correlation is difficult. However, vegetation development at the time of the QM zone at both sites was similar; it is especially characterized by the facts that Ulmus clearly migrated to the site earlier than Quercus and was very abundant throughout this time. Furthermore, both diagrams show very low amounts of Corylus. The interglacial of the Osterholz shows in addition to the above; a Carpinus-QM-Picea-zone in which Eucommia reaches a relative high value and in the upper of which Azolla filiculoides was also found. The similarity of vegetation development justifies acceptance of the same age for the occurrences. A comparison of the vegetation development at the Elm and the Osterholz with those of the Eem, Holstein, Waal, and Tegelen warm periods as well as with all the Cromer sites so far investigated shows that only a correlation with the Cromer Complex is possible. This correlation is supported by the geologic relations in the Osterholz (the deposit is overlain by Elster till). Therefore the till-like material with Scandinavian rock fragments underlying the deposit at Elm is of particular interest. The 'Rhume' interglacial beds at Bilshausen, only 60 km south of Osterholz, is also assigned to the Cromer complex, but the two deposits cannot be of the same age because the vegetation development differs. Therefore the Cromer complex must include at least two warm periods. Further conclusions about the relative stratigraphic position of these two occurrences and correlations of other Cromer sites are at this time not possible, however.

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The distribution of pollen in marine sediments is used to reconstruct pathways of terrigenous input to the oceans and provides a record of vegetation change on adjacent continents. The wind transport routes of aeolian pollen is comprehensively illustrated by clusters of trajectories. Isobaric, 4-day backward trajectories are calculated using the modelled wind-field of ECHAM3, and are clustered on a seasonal basis to estimate the main pathways of aeolian particles to sites of marine cores in the south-eastern Atlantic. Trajectories and clusters based on the modelled wind-field of the Last Glacial Maximum hardly differ from those of the present-day. Trajectory clusters show three regional, and two seasonal patterns, determining the pathways of aeolian pollen transport into the south-eastern Atlantic ocean. Mainly, transport out of the continent occurs during austral fall and winter, when easterly and south-easterly winds prevail. South of 25°S, winds blow mostly from the west and southwest, and aeolian terrestrial input is very low. Generally, a good latitudinal correspondence exists between the distribution patterns of pollen in marine surface sediments and the occurrence of the source plants on the adjacent continent. The northern Angola Basin receives pollen and spores from the Congolian and Zambezian forests mainly through river discharge. The Zambezian vegetation zone is the main source area for wind-blown pollen in sediments of the Angola Basin, while the semi-desert and desert areas are the main sources for pollen in sediments of the Walvis Basin and on the Walvis Ridge. A transect of six marine pollen records along the south-western African coast indicates considerable changes in the vegetation of southern Africa between glacial and interglacial periods. Important changes in the vegetation are the decline of forests in equatorial Africa and the north of southern Africa and a northward shift of winter rain vegetation along the western escarpment.