泽米科植物羽片脉序和解剖学及其系统学意义

研究了苏铁目泽米科Zamiaceae 2亚科的所有4族(Stevenson系统, 1992)共10种代表植物的羽片脉序及解剖学特征,结果显示泽米科羽片脉序为二歧分叉的平行脉,无中脉。小刺双子铁Dioon spinulosum、大头非洲铁Encephalartos friderici-guilielmii和摩尔大泽米Macrozamia moorei等的平行脉末端以不同的形式互相连接,而鳞木铁Lepidozamia peroffskyana、粗壮角果铁Ceratozamia mexicana var. robusta、竹叶角果铁C. hildae、佛州泽米Zamia　floridana、柔叶泽米Z. debilis、鳞秕泽米Z. furfuracea和短尖泽米Z. muricata等的平行脉末端不连接而直达叶缘,其中鳞木铁、粗壮角果铁和竹叶角果铁的脉达叶缘后逐渐消失。羽片的横切面结构通常由表皮、下皮厚壁细胞和叶肉组成,表皮层包括上、下表皮各一层,叶肉可能同时分化出近上表面的栅栏组织和近下表面的栅栏组织,或仅有近上表面的栅栏组织分化,或无栅栏组织分化而完全为海绵组织。然而,泽米科没有典型的海绵组织和传输组织分化。小刺双子铁、大头非洲铁、鳞叶木铁和摩尔大泽米的羽片具有粘液道而无工字厚壁组织,在小刺双子铁中粘液道与维管束对生,在另3种中则与维管束轮生;但粗壮角果铁、竹叶角果铁、佛州泽米、柔叶泽米、鳞秕泽米和短尖泽米的羽片则具有工字厚壁组织而没有粘液道,其中粗壮角果铁和竹叶角果铁的羽片工字厚壁组织仅与上表皮相连,而佛州泽米、柔叶泽米、鳞秕泽米和短尖泽米的羽片工字厚壁组织与上、下表皮都相连。羽片脉序和解剖学特征支持Stevenson将泽米铁科分为两亚科的观点。     

贺氏双子铁与摩瑞大泽米羽片和叶轴的解剖学研究

对贺氏双子铁和摩瑞大泽米的羽片和叶轴进行了比较研究,对两种植物的结构特征与对环境的适应性进行了分析,结果表明：两个种类的结构有很大的差异。贺氏双子铁的羽片仅在下表皮有气孔器分布,在近轴面仅有1～2层较短的栅栏组织,维管束与粘液道在垂周方向相对而生;摩瑞大泽米羽片在上下表皮均有气孔器,近轴面和远轴面均有较长的2～3层的栅栏组织,为等面叶,维管束与粘液道在平周方向相间排列。此外,两个种的维管束结构及叶缘结构等也有很大的差异。在叶轴方面,贺氏双子铁的表皮气孔数比摩瑞大泽米的气孔数明显要少,而且摩瑞大泽米的表皮内侧有一环富含叶绿体的栅栏组织,且在栅栏组织的内方还有发达的富含叶绿体的海绵组织,而贺氏双子铁则没有。此外,摩瑞大泽米在基本组织中含有较多的副转输组织,而贺氏双子铁则无。在粘液道大小、排列方式上两种类也有很大的不同。贺氏双子铁的维管束韧皮部明显比木质部发达,而摩瑞大泽米木质部和韧皮部的发达程度相近。     

三种苏铁植物呈现出迥异的水力安全边界北大核心CSCD

为探究苏铁植物的水力安全边界(hydraulic safety margins,HSM),该试验选用经典的自然干燥法和最新发表的抽气法测定三种同质园苏铁植物抗旱性(即叶轴木质部脆弱性曲线),获得抗旱指标P_(50)和P_(88)(导水率丧失或气体抽取量分别为50%和88%时的木质部水势),与叶片正午水势计算HSM,结合管胞性状分析。结果表明:(1)苏铁(Cycas revoluta)、越南篦齿苏铁(C.elongata)、摩瑞大泽米苏铁(Macrozamia moorei)的管胞长度分别为(4413±378)、(5146±730)、(6954±862)μm,苏铁、越南篦齿苏铁与摩瑞大泽米苏铁差异显著(P<0.05)。(2)两种方法测定的脆弱性曲线都呈典型的“S”型,苏铁、越南篦齿苏铁、摩瑞大泽米苏铁的P_(50H)(导水率丧失50%时的木质部水势)和P_(50P)(气体抽取量为50%时的木质部水势)分别为-2.5、(-2.4±0.5)MPa,-2.3、(-3.6±0.7)MPa,-1.5、(-1.8±0.2)MPa,在已发表的裸子植物数值范围内。P_(50)和P_(88)具有显著一致性(除了越南篦齿苏铁的P_(50P)比P_(50H)更低,表示更强抗旱性),与已发表的其他木质部管胞物种通过水力学法和抽气法获得的P_(50)和P_(88)比较分析,具有显著相关性(R^(2)=0.72,P=0.0081;R^(2)=0.87,P=0.0007)。(3)自然干燥法和抽气法计算的HSMs具有相同的趋势,摩瑞大泽米苏铁为负值,而苏铁和越南篦齿苏铁为正值。综上所述,三种苏铁植物的抗旱性均在已发表的裸子植物范围内,两种方法都适于测定木质部管胞结构的苏铁类植物脆弱性曲线,苏铁、越南篦齿苏铁与摩瑞大泽米苏铁具有不同的水力安全边界。利用脆弱性曲线和正午水势探讨苏铁植物的水力安全边界,为苏铁植物的水分监测、管理和保育提供依据。     

托叶铁科(苏铁目)羽片脉序和羽片解剖学研究

详细研究了托叶铁科两属的2种代表植物Stangeria eriopus和Bowenia serrulata的羽片脉序和羽片解剖学,两个属在气孔器特征与不具副传输组织方面极为相似。而在气孔的分布,羽片脉序式样与叶缘形态,粘液道的有无,海绵组织与栅栏组织的分化。工字厚壁组织与表皮细胞垂周壁特征方面则有较大的差异。将这些特征与苏铁科和泽米铁科作了比较,并讨论了这些特征在系统学上的意义。研究结果支持Stevenson(1992)将该科分为两亚科以及托叶铁科是介入苏铁科与泽米铁科之间的观点。     

越南篦齿苏铁传粉媒介的研究

利用排除法和引入昆虫的方法相结合,研究越南篦齿苏铁(Cycas elongata)的传粉媒介.结果表明,越南篦齿苏铁靠风媒传粉和虫媒传粉的雌株结实率分别55.3%和57.2%,自然传粉的结实率为63.5%,说明风和象鼻虫都是越南篦齿苏铁的有效传粉媒介.越南篦齿苏铁散粉高峰在白天,夜间散粉很少,在3.0 m以内风传花粉的密度较高,3.0 m之外的密度急剧下降.可见,风媒和虫媒都是越南篦齿苏铁的有效传粉途径,与泽米铁类植物为专一寄主性昆虫传粉的结论不一致.     

美洲来的铁树——鳞秕泽米

鳞秕泽米(Zamia furfuracea)为近年从中美洲引进的苏铁科(广义的)或泽米科(狭义的)泽米属观叶植物。其叶片、株形较之于苏铁更多一种粗犷豪放的精神,在国内日益受到花卉爱好者的青睐。 鳞秕泽米树干多半生于地下,多分枝,干高15—30厘米,圆柱状,树皮密布暗褐色鳞秕状叶痕,多年生的总干基部茎盘处常着生幼小的萌蘖。叶为大型偶数羽状复叶,8—25片丛生     

岩蕨科、球盖蕨科的系统位置：叶绿体DNAtrnL—F区序列证据

岩蕨科（Woodsiaceae）和球盖蕨科（Peranemaceae）历来被多数学者认为是亲缘关系密切的两个科,它们的系统位置一直存在争议,两科曾被放入三叉蕨科（Aspidiaceae）、碗蕨科（Dennstaedtiaceae）、鳞毛蕨亚科（Dryopteridoidaae）及鳞毛蕨科（Dryopteriaceae）蹄盖蕨亚科（Athyrioideae）中。根据岩蕨科1种、球盖蕨科3种、蹄盖蕨科8种、鳞毛蕨科2种及1种外类群假大羽铁角蕨（Asplenium pseudolaserpitiifolium）的cpDNA基因trnL—F区序列建立的系统树,结果显示,岩蕨科和球盖蕨科的亲缘关系较远,岩蕨科与蹄盖蕨科有密切的亲缘关系,球盖蕨科与鳞毛蕨科亲缘关系较近,岩蕨科和球盖蕨科是处于不同进化线上的植物。秦仁昌（1978）和Pichi—Sermolii（1977）的系统将二者分别作为独立的科是合适的。     

泽米铁科三种植物的叶表皮特征

在光镜下观察了泽米铁科分属于3个属的3种植物叶表皮特征,结果表明：3种植物叶表皮细胞及气孔器差异显著,一定程度上证实了3属是自然的分类群。在鳞秕泽米铁(Zamia furfuracea)中发现了气孔簇。本文还对3种植物之间的系统演化关系及它们对环境的适应性进行了讨论。     

贵州苏铁的核型分析

本文报道了贵州苏铁(Cycas guizhouensis)的染色体数目及核型,并与攀枝花苏铁(C.panzhihuaensis)、苏铁(C.revoluta)、华南苏铁(C.rumphii),拳叶苏铁(C.circinalis)和C.media等的核型进行了比较,讨论了苏铁属植物不同核型的进化趋势,提出了苏铁属植物的两种地理分布核型,即1.北回归线以北为K(2n)=22=2m+4sm+4st+12T包括贵州苏铁和攀枝花苏铁;2.北回归线及其以南为K(2n)=22=4m+8st+10T,包括苏铁、华南苏铁、拳叶苏铁和C.media。     

岩蕨科、球盖蕨科的系统位置：叶绿体DNAtrnL-F区序列证据

岩蕨科(Woodsiaceae)和球盖蕨科(Peranemaceae)历来被多数学者认为是亲缘关系密切的两个科,它们的系统位置一直存在争议,两科曾被放入三叉蕨科(Aspidiaceae)、碗蕨科(Dennstaedtiaceae)、鳞毛蕨亚科(Dryopteridoidaae)及鳞毛蕨科(Dryopteriaceae)蹄盖蕨亚科(Athyrioideae)中。根据岩蕨科1种、球盖蕨科3种、蹄盖蕨科8种、鳞毛蕨科2种及1种外类群假大羽铁角蕨(Asplenium pseudolaserpitiifolium)的cpDNA基因trnL-F区序列建立的系统树,结果显示,岩蕨科和球盖蕨科的亲缘关系较远,岩蕨科与蹄盖蕨科有密切的亲缘关系,球盖蕨科与鳞毛蕨科亲缘关系较近,岩蕨科和球盖蕨科是处于不同进化线上的植物。秦仁昌（1978）和Pichi-Sermolii（1977）的系统将二者分别作为独立的科是合适的。     

福建苏铁调查研究初报

对福建苏铁经过三年多的调查、观察和研究 ,基本完成福建苏铁原产地的调查 ,认为福建原产两种苏铁 ,即苏铁与四川苏铁。本文还对福州树木园引种栽培的苏铁进行分类研究 ,有苏铁、四川苏铁、海南苏铁、攀枝花苏铁、广东苏铁、台东苏铁、元江苏铁、暹罗苏铁、石山苏铁、篦齿苏铁、越南苏铁、叉叶苏铁、多羽叉叶苏铁、多歧苏铁、刺叶苏铁等 ,另有三种形态特征比较特殊的苏铁也在本文进行了简要描述。     

福建苏铁调查研究初报

对福建苏铁经过三年多的调查、观察和研究,基本完成福建苏铁原产地的调查,认为福建原产两种苏铁,即苏铁与四川苏铁。本文还对福州树木园引种栽培的苏铁进行分类研究,有苏铁、四川苏铁、海南苏铁、攀枝花苏铁、广东苏铁、台东苏铁、元江苏铁、暹罗苏铁、石山苏铁、篦齿苏铁、越南苏铁、叉叶苏铁、多羽叉叶苏铁、多歧苏铁、刺叶苏铁等,另有三种形态特征比较特殊的苏铁也在本文进行了简要描述。     

基于RAPD分析的中国苏铁属部分种类亲缘关系探讨

利用21个筛选出来的RAPD引物,对苏铁属21个种的22份材料进行分析,获得333个RAPD标记,利用NTSYS(V.2.10e)软件,建立了22份供试材料的UPGMA聚类图,进而探讨了苏铁属21个种类间的亲缘关系。RAPD聚类分析结合形态学研究结果表明:多裂苏铁和叉孢苏铁的亲缘关系很近,聚为一类,多裂苏铁应为叉孢苏铁的一个亚种。西林苏铁、隆林苏铁、叉孢苏铁、尖尾苏铁、叉叶苏铁、长柄叉叶苏铁、多羽叉叶苏铁、长球果苏铁、贵州苏铁、四川苏铁、短叶苏铁、石山苏铁、宽叶苏铁、十万大山苏铁、元江苏铁、仙湖苏铁、海南苏铁、台湾苏铁、广东苏铁、滇南苏铁相互间的亲缘关系均较远,支持各自为独立的种。     

A phylogeny of cycads (Cycadales) inferred from chloroplast matK gene, trnK intron, and nuclear rDNA ITS region

Phylogenetic relationships among the three families and 12 living genera of cycads were reconstructed by distance and parsimony criteria using three markers: the chloroplast matK gene, the chloroplast trnK intron and the nuclear ITS/5.8S rDNA sequence. All datasets indicate that Cycadaceae (including only the genus Cycas) is remotely related to other cycads, in which Dioon was resolved as the basal-most clade, followed by Bowenia and a clade containing the remaining nine genera. Encephalartos and Lepidozamia are closer to each other than to Macrozamia. The African genus Stangeria is embedded within the New World subfamily Zamiodeae. Therefore, Bowenia is an unlikely sister to Stangeria, contrary to the view that they form the Stangeriaceae. The generic status of Dyerocycas and Chigua is unsupportable as they are paraphyletic with Cycas and the Zamia, respectively. Nonsense mutations in the matK gene and indels in the other two datasets lend evidence to reinforce the above conclusions. According to the phylogenies, the past geography of the genera of cycads and the evolution of character states are hypothesized and discussed. Within the suborder Zamiieae, Stangeria, and the tribe Zamieae evolved significantly faster than other genera. The matK gene and ITS/5.8S region contain more useful information than the trnK intron in addressing phylogeny. Redelimitations of Zamiaceae, Stangeriaceae, subfamily Encephalartoideae and subtribe Macrozamiineae are necessary.     

苏铁属植物RAPD反应体系的优化及部分种类亲缘关系的探索

通过优化苏铁属植物的RAPD反应体系,进而探讨苏铁属部分种类的亲缘关系。结果表明,Mg~(2+)、dNTP、Taq酶及随机引物浓度在RAPD反应中有重要影响,而模板DNA浓度有一个很大的适应范围。适合苏铁属植物RAPD分析的反应体系:20μL反应体系中,含有10 mmol/L Tris-HCl(pH8.3)、50 mmol/L KCl、2.0mmol/L MgCl_2、200μmol/L dNTP、0.3μmol/L随机引物、模板DNA 50ng、Taq酶1.0 U。聚类分析结果基本反应了苏铁属各个种间的亲缘关系,证明了RAPD适用于苏铁属种间亲缘关系分析。RAPD聚类分析结合形态学研究表明:海南苏铁、台湾苏铁、广东苏铁、滇南苏铁和仙湖苏铁之间的亲缘关系较远,支持各自成为独立的种。     

A new species of Leptocycas(Zamiaceae)from the Upper Triassic sediments of Liaoning Province,China

A new cycad,Leptocycas yangcaogouensis sp.nov.,was found in sediments from the Late Triassic in western Liaoning,China.The pinnately compound leaves(Pseudoctenis type)are screwed in a crown on the stem top.The leaflets are linear,with parallel veins and decurrent bases on the rachis.The leaf bases are persistent.The cataphylls intermix with the leaves.The female cone is ovoid in shape.The characteristics of the new plant are more similar to those of Leptocycas gracilis,a Triassic cycad from North America,but the new species differs from L.gracilis in the size of its stem(7-8 vs.3-5 cm in diameter,respectively),leaves(length × width 100 × 16 vs.30 × 7 cm,respectively)and leaf density along the stem(4-6 vs.1-2 bases/1 cm length,respectively).Both L.gracilis and L.yangcaogouensis,having leaves of the Pseudoctenis type,show a closer relationship to the extant Dioon of Zamiaceae.The present study provides evidence for the origin of the genus Dioon,which may have come from Leptocycas plants of the Triassic.It would be assumed that the extent cycads in Zamiaceae originate from the pteridosperms in the Late Paleozoic and have evolved through the stage of L.gracilis and L.yangcaogouensis in Late Triassic,and reaching the extant Dioon.     

Is Cycas revoluta (Cycadaceae) wind- or insect-pollinated?

Among the Cycadales (Cycadaceae and Zamiaceae), the Zamiaceae are known to be insect-pollinated. In contrast, the Cycadaceae are still considered wind-pollinated, although some doubt has been cast on several species, including Cycas revoluta. Using a large population of C. revoluta on Yonaguni Island (Okinawa, Japan), we performed exclusion experiments, documented insects from male and female cones, and analyzed the morphology of the apical part of the ovule to determine the pollination method of this species. Insect exclusion resulted in a notable reduction in seed set, except in a few individuals growing near male cones. The amount of airborne pollen was abundant within a 2-m radius of male cones but decreased markedly beyond this distance. Pollen grains of C. revoluta were found on the body of Carpophilus chalybeus (Nitidulidae, Coleoptera), one of a few species of insects collected from both male cones and female cones far from males. We conclude that C. revoluta relies on both wind (anemophily) and insect pollination (entomophily), although such anemophily is restricted to female trees growing within a 2-m radius of male trees. The nitidulids are not host specific to this cycad and primarily feed on plant tissue but serve as pollinators during pollen release. Cycas revoluta appears to be in an initial mode of animal pollination, as opposed to the host-specific insect pollination observed in most Zamiaceae.     

A phylogenetic analysis of American Zamiaceae (Cycadales) using chloroplast DNA restriction fragment length polymorphisms

A phylogenetic analysis of American cycad genera, all belonging to the family Zamiaceae, was attempted using chloroplast DNA restriction fragment polymorphisms.Ceratozamia mexicana Brongn.,Chigua restrepoi D. Stevenson,Dioon edule Lindley,Microcycas calocoma (Miq.) A. DC.,Zamia fischeri Miq., andZamia skinneri Warsz. ex A. Dietrich were used as representatives of the genera.Cycas revoluta Thunb., belonging to the family Cycadaceae, was used as an outgroup, following previous morphological works. One hundred and forty-one shared restriction fragments were scored for presence/absence and both Wagner and Dollo parsimony analyses were performed. The single, fully resolved, most parsimonious trees obtained from the analyses were topologically identical and perfectly matched previous morphology-based phylogenetic hypotheses. Statistical evaluation of the data showed a good reliability for the obtained phylogeny.Dioon edule, belonging to a different subfamily and more primitive on morphological grounds, proved to be the most primitive among American cycads as inferred from the molecular data;Chigua restrepoi, never analyzed before on cladistic grounds, was found to be the sister group of the genusZamia.