伞形科部分类群气孔结构及其分类学价值

郭诗琦，王超群，刘　玫，茹　剑，王靖茹，王雪微

(哈尔滨师范大学 生命科学与技术学院，黑龙江省普通高等学校植物生物学重点实验室，哈尔滨 150025)



伞形科部分类群气孔结构及其分类学价值

郭诗琦，王超群，刘玫*，茹剑，王靖茹，王雪微

(哈尔滨师范大学 生命科学与技术学院，黑龙江省普通高等学校植物生物学重点实验室，哈尔滨 150025)

该研究对变豆菜亚科及芹亚科基础类群20属29种(变豆菜亚科9属11种，芹亚科基础类群11属18种)植物叶片(苞片或果实)的气孔结构进行显微观察分析，以明确变豆菜亚科及芹亚科基础类群的气孔特征及其与伞形科其他类群的区别。结果显示：(1)变豆菜亚科及芹亚科基础类群的气孔类型有无规则型、不等型和平列型3种，其中变豆菜亚科的变豆菜族气孔为无规则型(占总气孔比例的35%～75%)及不等型(25%～65%)，仅刺芹属(Eryngium)具平列型，而Steganotaenieae族的气孔均为无规则型；芹亚科基础类群除柴胡属(Bupleurum)外多为或仅为无规则型(75%～100%)，少为不等型(5%～25%)。(2)气孔结构支持分子系统学将Arctopus放入变豆菜亚科的变豆菜族，Choritaenia移入芹亚科。(3)变豆菜亚科气孔特征同芹亚科及牵环花亚科的较为相似，而同参棕亚科差别较大。

伞形科；气孔；变豆菜亚科；芹亚科；系统学

Drude根据果实特征将伞形科(Apiaceae)分为三亚科，天胡荽亚科(Hydrocotyloideae)，变豆菜亚科(Saniculoideae)及芹亚科(Apioideae)[1]。分子系统学研究显示天胡荽亚科，不是一个自然的类群[2-8]，其中大部分属应分别位于2个新亚科，参棕亚科(Mackinlaloideae)和牵环花亚科(Azorelloideae)[2]。变豆菜亚科主要分布在北半球，植物具单叶，头状花序，叶片边缘锯齿状或多刺，果实表面具刺、薄片或瘤状突起，内果皮为薄壁细胞构成，油管主要分布在主棱。此亚科包括2个族：变豆菜族(Sanicluleae)[6属：Actinolema，Alepidea，Astrantia，刺芹属(Eryngium)，Hacquetia及变豆菜属(Sanicula)]和Lagoecieae族(2属：Lagoecia及Petagnaea)[1,9]。非洲特有属Arctopus花的许多特征存于天胡荽亚科[10]，曾被位于天胡荽亚科不确定类群[9]，2个木本属Polemanniopsis和Steganotaenia原位于芹亚科不确定类群及Tordylieae族[9]，然而分子系统学研究显示这3属靠近变豆菜亚科[2,7,11]，Lagoecia属移出变豆菜亚科，位于芹亚科并靠近海茴香(Crithmummaritimum)[8,12-13]。随着分子系统学研究的发展，Calvio和Downie[14]指出变豆菜亚科包括2个族：变豆菜族(Saniculeae)及Steganotaenieae，前者包括Actinolema，Alepidea，Arctopus，Astrantia，刺芹属(Eryngium)及Petagnaea，后者包括Polemanniopsis和Steganotaenia。

与变豆菜亚科不同，芹亚科多数物种具有伞形花序，游离的心皮柄，发达的油管，然而此亚科不是自然的类群[15]。一些非洲特有属(原位于Apieae族[9])是芹亚科的基础类群，其中5属(Dracosciadium，Anginon，Glia，Heteromorpha及Polemannia)组成一个新的族(Heteromorpheae)，同Annesorhizaclade(Annesorhiza及Chamarea)，Phlyctidocarpa，Lichtensteinia及Choritaenia属的亲缘关系较近。除此之外，芹亚科基础类群还包括柴胡属(Bupleurum)[3,7,12-13,15-17]。

尽管已有研究揭示了伞形科果实微形态对其系统学研究的价值[18-19]，但有许多研究表明气孔类型具有分类学价值。变豆菜亚科刺芹属(Eryngium)的某些种(如E.reticum及E.compestre)具横列型气孔(diacytic)[20]，芹亚科具有不同类型的气孔，如无规则型(anomocytic)、不等型(anisocytic)、横列型及平列型(paracytic)[21-25]。然而缺少对于整个变豆菜亚科及芹亚科基础类群气孔结构的研究。本研究旨在深入研究变豆菜亚科及芹亚科基础类群的气孔特征，掌握他们气孔特征及与伞形科其他类群的区别，弄清气孔结构是否可为分子系统学的研究提供形态学依据。

1　材料和方法

1.1材料

本研究材料为变豆菜亚科(9属11种)及芹亚科基础类群(11属18种)，共20属29种植物的叶片(极少数物种因未得到叶片而用其苞片或果实)。材料分别取自于中国科学院西双版纳热带植物园标本馆(HITBC-Xishuangbanna Tropical Botanical Garden Herbarium, CAS)，中国科学院华南植物园标本馆(IBSC-South China Botanical Garden Herbarium)，南非约堡大学植物及植物生物技术系标本馆(JRAU-Herbarium, Department of Botany and Plant Biotechnology, University of Johannesburg, South Africa)，俄罗斯莫斯科州立大学生物学院标本馆(MW-Herbariun, Biological Faculty, Moscov State University, Moscov, Russia)，南非康普顿标本馆(NBG-Compton Herbarium of South Africa)，美国纽约植物园标本馆(NY-New York Botanical Garden Herbarium, America)，中国科学院植物研究所标本馆(PE-Herbarium, Institute of Botany, CAS)，南非国家标本馆(PRE-National Herbarium of South Africa)，纳米比亚农业和自然保护部西南非洲植物标本馆(WIND-South-West Africa Herbarium, Department of Agriculture and National Conseration, Namibia)。物种名称及凭证标本信息见表1 (其中仅软雀花Saniculaelata和黑柴胡Bupleurumsmithii为中国产物种)。

1.2方法

将叶片(苞片或果实)浸泡在约90℃热水中，待材料充分吸水(约15～30 min)后将材料取出，根据Zarinkamar[26]的观察(伞形科植物叶片的气孔密度，下表皮大于上表皮)，撕取植物叶片(或苞片)的下表皮(或果皮)，将其放在载玻片上，加1～2滴50%甘油水溶液并加盖玻片。用具成像系统(Olympus DP 70)的显微镜(Olympus BX 51)观察气孔结构并照相。每个样品观察100个气孔并统计计算不同类型气孔数量及所占比例，根据照片用Photoshop CS2.0软件绘制气孔图。

2　结果与讨论

研究显示变豆菜亚科具有3种类型气孔：无规则型、不等型及平列型(图版Ⅰ，A～I)。其中多数物种具有无规则型(35%～75%)及不等型(25%～65%)。不等型多于无规则型(Actinolemaeryngioides、Alepideaamatymbica、A.longifolia、Arctopusechinatus及A.monacanthus，图版Ⅰ，A～C)或无规则型多于不等型(Astrantiatrifida，Hacquetiaepipactis及Saniculaelata，图版Ⅰ，D、F、H)。Polemanniopsismarlothii及Steganotaeniaaraliacea仅具有无规则型(图版Ⅰ，G、I)，Eryngiumamethystinum(图版Ⅰ，E)具3种类型气孔：无规则型(30%)，不等型(30%)及平列型(40%)。芹亚科基础类群仅具无规则型及不等型(图版Ⅱ，A～R)。Annesorhizaflagellifolia，A.hirsute及Gliapilulosa(图版Ⅱ，B～C、K)的气孔多为无规则型(75%～100%)，少为不等型(5%～25%)。Bupleurumsmithii(图版Ⅱ，D)的不等型气孔(80%)多于无规则型(20%)。所研究物种的表皮细胞垂周壁通常是直的，但少数物种的细胞壁波状弯曲，如Astrantiatrifida、Polemanniopsismarlothii及Phlyctidocarpaflava(图版Ⅰ，D、G；图版Ⅱ，O)，且同一属(如Chamarea)的不同物种(如C.esterhuyseniae及C.gracillima)的细胞壁可以是平直或弯曲(图版Ⅱ，E、F)。

表1　用于研究气孔结构的伞形科变豆菜亚科及芹亚科基础类群的凭证标本信息及其采集地Table 1　Taxa of Apiaceae subfamilies Saniculoideae and basal Apioideae examined for stomatal characters, together with voucher specimen details and locality

分子系统学将Polemanniopsis及Steganotaenia两属从芹亚科移入变豆菜亚科[6-7,14]，果实微形态显示二者果皮中有大量草酸钙结晶及脊上有大油管，同变豆菜亚科物种很相似[27]，但两属依然有芹亚科物种具有的特点，如复伞形花序的特点普遍存在于芹亚科，果实具侧脊形成翅存在于芹亚科某些类群(如Heteromorpha)。本研究揭示Polemanniopsis及Steganotaenia具有芹亚科的气孔特征(表2)。变豆菜亚科的变豆菜族主要有无规则型及不等型气孔可以区别于Steganotaenieae族。刺芹属的平列型气孔主要存在于参棕亚科，而牵环花亚科的气孔与本研究揭示的气孔类型相似，主要为无规则型及不等型。其他刺芹属物种的气孔类型需要进一步研究，从而探讨此属同变豆菜族其他类群的关系。

芹亚科植物通常为复叶，但柴胡属具单叶[28-30]，单叶存在于参棕亚科，牵环花亚科及变豆菜亚科。此外柴胡属花粉特征也同这些类群相似，为近菱形，近圆形及椭圆形[31-32]。分子系统研究证明柴胡属是单系类群，比非洲特有属进化，但仍位于芹亚科的基础位置[5,7,33]。果实微形态显示所研究的来自非洲的芹亚科基础类群(Annesorhizaclade，Heteromorpheae，Lichtensteiniaclade，Phlyctidocarpa)的果皮均分布草酸钙结晶，根据Burtt[34]的描述，芹亚科物种通常没有结晶，如果有结晶仅存在果实的结合面。本研究显示除柴胡属外芹亚科基础类群的气孔主要或仅为无规则型。而柴胡属属内物种气孔结构的变化有待于进一步探究。Choritaenia原位于天胡荽亚科[9]，果实具特殊的油囊[35]，以及气孔结构均显示此属与芹亚科物种相似，支持分子系统学将其移入芹亚科。

表2　伞形科变豆菜亚科及芹亚科基础类群叶片(苞片或果实)表皮气孔类型Table 2　Summary of stomatal types of leaf (bract or fruit) epidermis in Apiaceae subfamilies Saniculoideae and basal Apioideae taxa

注：*苞片，**果实

Note: *bract，**fruit

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图版Ⅰ　伞形科变豆菜亚科植物叶(苞片或果实)表皮气孔结构(标尺=20 μm)ano.无规则型气孔; ani.不等型气孔; pa.平列型气孔Fig. A. Actinolema eryngioides; Fig. B. Alepidea amatymbica; Fig. C. Arctopus monacanthus; Fig. D. Astrantia trifida; Fig. E. Eryngium amethystinum; Fig. F. Hacquetia epipactis; Fig. G. Polemanniopsis marlothii; Fig. H. Sanicula elata; Fig. I. Steganotaenia araliaceaPlateⅠ　The stomatal structures of the leaf (bract or fruit) epidermis of the taxa in subfamily Saniculoideae. (Scale bar=20 μm)ano. Anomocytic; ani. Anisocytic; pa. Paracytic

图版Ⅱ　伞形科芹亚科基础类群叶(或果实)表皮气孔结构(标尺=20 μm)ano.无规则型气孔；ani.不等型气孔Fig.A. Anginon fruticosum；Fig. B. Annesorhiza flagellifolia；Fig.C. A. hirsute；Fig.D. Bupleurum smithii；Fig.E. Chamarea esterhuyseniae；Fig.F. C. Gracillima；Fig.G. Choritaenia capensis；Fig.H. Dracosciadium italae；Fig.I. D. Saniculifolium；Fig.J. Glia gummifera；Fig.K. G. pilulosa；Fig. L. Heteromorpha involucrata；Fig.M. H. stolzii；Fig. N. Polemannia grossulariifolia；Fig.O. Phlyctidocarpa flava；Fig.P. P. montana; Fig.Q. Lichtensteinia lacera；Fig.R. L. trifidaPlateⅡ　The stomatal structures of the leaf (or fruit) epidermis in basal Apioideae taxa (Scale bar =20 μm)ano. Anomocytic；ani. Anisocytic

(编辑:潘新社)

Stomatal Structures of Some Taxa in Apiaceae and Their Taxonomic Values

GUO Shiqi, WANG Chaoqun, LIU Mei*, RU Jian, WANG Jingru, WANG Xuewei

(College of Life Science and Technology, Harbin Normal University, Key laboratory of Plant Biology, College of Heilongjiang Province, Harbin 150025, China)

The stomatal structures of leaves (bracts or fruits) from 20 genera and 29 species (9 genera and 11 species of subfamily Saniculoideae, 11 genera and 18 species of basal subfamily Apioideae). The results show that: (1) three types of stomata, anomocytic, anisocytic, and paracytic. The stomata in tribe Saniculeae of Saniculoideae are anomocytic (35%-75%) and anisocytic (25%-65%) (in the total number of stomata). The genusEryngiumhas all three types, and only anomocytic type in tribe Steganotaenieae. In the basal taxa of subfamily Apioideae (exceptBupleurum) the stomata are more anomocytic (75%-100%) and less anisocytic (5%-25%). (2)The stomatal features studied support to placeArctopusin Saniculeae and to moveChoritaeniato Apioideae confirmed by molecular data. (3) The stomata structures of Saniculoideae are similar with those of Apioideae and Azorelloideae, but differ from Mackinlayoideae.

Apiaceae; stoma; Saniculoideae; Apioideae; phylogeny

1000-4025(2016)09-1787-07doi:10.7606/j.issn.1000-4025.2016.09.1787

2016-04-07;修改稿收到日期:2016-09-01

国家自然科学基金(31270235,31070169)

郭诗琦(1994-)，女，学士，主要从事植物学研究。E-mail:guosq17@163.com

刘玫，教授，博士导师，主要从事植物学科研及教学工作。E-mail:m.r.liu@126.com

Q246; Q944.53

A