傘形科部分類群氣孔結構及其分類學價值

郭詩琦，王超群，劉　玫，茹　劍，王靖茹，王雪微

(哈爾濱師范大學 生命科學與技術學院，黑龍江省普通高等學校植物生物學重點實驗室，哈爾濱 150025)

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傘形科部分類群氣孔結構及其分類學價值

郭詩琦，王超群，劉玫*，茹劍，王靖茹，王雪微

(哈爾濱師范大學 生命科學與技術學院，黑龍江省普通高等學校植物生物學重點實驗室，哈爾濱 150025)

該研究對變豆菜亞科及芹亞科基礎類群20屬29種(變豆菜亞科9屬11種，芹亞科基礎類群11屬18種)植物葉片(苞片或果實)的氣孔結構進行顯微觀察分析，以明確變豆菜亞科及芹亞科基礎類群的氣孔特征及其與傘形科其他類群的區(qū)別。結果顯示：(1)變豆菜亞科及芹亞科基礎類群的氣孔類型有無規(guī)則型、不等型和平列型3種，其中變豆菜亞科的變豆菜族氣孔為無規(guī)則型(占總氣孔比例的35%～75%)及不等型(25%～65%)，僅刺芹屬(Eryngium)具平列型，而Steganotaenieae族的氣孔均為無規(guī)則型；芹亞科基礎類群除柴胡屬(Bupleurum)外多為或僅為無規(guī)則型(75%～100%)，少為不等型(5%～25%)。(2)氣孔結構支持分子系統(tǒng)學將Arctopus放入變豆菜亞科的變豆菜族，Choritaenia移入芹亞科。(3)變豆菜亞科氣孔特征同芹亞科及牽環(huán)花亞科的較為相似，而同參棕亞科差別較大。

傘形科；氣孔；變豆菜亞科；芹亞科；系統(tǒng)學

Drude根據果實特征將傘形科(Apiaceae)分為三亞科，天胡荽亞科(Hydrocotyloideae)，變豆菜亞科(Saniculoideae)及芹亞科(Apioideae)[1]。分子系統(tǒng)學研究顯示天胡荽亞科，不是一個自然的類群[2-8]，其中大部分屬應分別位于2個新亞科，參棕亞科(Mackinlaloideae)和牽環(huán)花亞科(Azorelloideae)[2]。變豆菜亞科主要分布在北半球，植物具單葉，頭狀花序，葉片邊緣鋸齒狀或多刺，果實表面具刺、薄片或瘤狀突起，內果皮為薄壁細胞構成，油管主要分布在主棱。此亞科包括2個族：變豆菜族(Sanicluleae)[6屬：Actinolema，Alepidea，Astrantia，刺芹屬(Eryngium)，Hacquetia及變豆菜屬(Sanicula)]和Lagoecieae族(2屬：Lagoecia及Petagnaea)[1,9]。非洲特有屬Arctopus花的許多特征存于天胡荽亞科[10]，曾被位于天胡荽亞科不確定類群[9]，2個木本屬Polemanniopsis和Steganotaenia原位于芹亞科不確定類群及Tordylieae族[9]，然而分子系統(tǒng)學研究顯示這3屬靠近變豆菜亞科[2,7,11]，Lagoecia屬移出變豆菜亞科，位于芹亞科并靠近海茴香(Crithmummaritimum)[8,12-13]。隨著分子系統(tǒng)學研究的發(fā)展，Calvio和Downie[14]指出變豆菜亞科包括2個族：變豆菜族(Saniculeae)及Steganotaenieae，前者包括Actinolema，Alepidea，Arctopus，Astrantia，刺芹屬(Eryngium)及Petagnaea，后者包括Polemanniopsis和Steganotaenia。

與變豆菜亞科不同，芹亞科多數物種具有傘形花序，游離的心皮柄，發(fā)達的油管，然而此亞科不是自然的類群[15]。一些非洲特有屬(原位于Apieae族[9])是芹亞科的基礎類群，其中5屬(Dracosciadium，Anginon，Glia，Heteromorpha及Polemannia)組成一個新的族(Heteromorpheae)，同Annesorhizaclade(Annesorhiza及Chamarea)，Phlyctidocarpa，Lichtensteinia及Choritaenia屬的親緣關系較近。除此之外，芹亞科基礎類群還包括柴胡屬(Bupleurum)[3,7,12-13,15-17]。

盡管已有研究揭示了傘形科果實微形態(tài)對其系統(tǒng)學研究的價值[18-19]，但有許多研究表明氣孔類型具有分類學價值。變豆菜亞科刺芹屬(Eryngium)的某些種(如E.reticum及E.compestre)具橫列型氣孔(diacytic)[20]，芹亞科具有不同類型的氣孔，如無規(guī)則型(anomocytic)、不等型(anisocytic)、橫列型及平列型(paracytic)[21-25]。然而缺少對于整個變豆菜亞科及芹亞科基礎類群氣孔結構的研究。本研究旨在深入研究變豆菜亞科及芹亞科基礎類群的氣孔特征，掌握他們氣孔特征及與傘形科其他類群的區(qū)別，弄清氣孔結構是否可為分子系統(tǒng)學的研究提供形態(tài)學依據。

1　材料和方法

1.1材料

本研究材料為變豆菜亞科(9屬11種)及芹亞科基礎類群(11屬18種)，共20屬29種植物的葉片(極少數物種因未得到葉片而用其苞片或果實)。材料分別取自于中國科學院西雙版納熱帶植物園標本館(HITBC-Xishuangbanna Tropical Botanical Garden Herbarium, CAS)，中國科學院華南植物園標本館(IBSC-South China Botanical Garden Herbarium)，南非約堡大學植物及植物生物技術系標本館(JRAU-Herbarium, Department of Botany and Plant Biotechnology, University of Johannesburg, South Africa)，俄羅斯莫斯科州立大學生物學院標本館(MW-Herbariun, Biological Faculty, Moscov State University, Moscov, Russia)，南非康普頓標本館(NBG-Compton Herbarium of South Africa)，美國紐約植物園標本館(NY-New York Botanical Garden Herbarium, America)，中國科學院植物研究所標本館(PE-Herbarium, Institute of Botany, CAS)，南非國家標本館(PRE-National Herbarium of South Africa)，納米比亞農業(yè)和自然保護部西南非洲植物標本館(WIND-South-West Africa Herbarium, Department of Agriculture and National Conseration, Namibia)。物種名稱及憑證標本信息見表1 (其中僅軟雀花Saniculaelata和黑柴胡Bupleurumsmithii為中國產物種)。

1.2方法

將葉片(苞片或果實)浸泡在約90℃熱水中，待材料充分吸水(約15～30 min)后將材料取出，根據Zarinkamar[26]的觀察(傘形科植物葉片的氣孔密度，下表皮大于上表皮)，撕取植物葉片(或苞片)的下表皮(或果皮)，將其放在載玻片上，加1～2滴50%甘油水溶液并加蓋玻片。用具成像系統(tǒng)(Olympus DP 70)的顯微鏡(Olympus BX 51)觀察氣孔結構并照相。每個樣品觀察100個氣孔并統(tǒng)計計算不同類型氣孔數量及所占比例，根據照片用Photoshop CS2.0軟件繪制氣孔圖。

2　結果與討論

研究顯示變豆菜亞科具有3種類型氣孔：無規(guī)則型、不等型及平列型(圖版Ⅰ，A～I)。其中多數物種具有無規(guī)則型(35%～75%)及不等型(25%～65%)。不等型多于無規(guī)則型(Actinolemaeryngioides、Alepideaamatymbica、A.longifolia、Arctopusechinatus及A.monacanthus，圖版Ⅰ，A～C)或無規(guī)則型多于不等型(Astrantiatrifida，Hacquetiaepipactis及Saniculaelata，圖版Ⅰ，D、F、H)。Polemanniopsismarlothii及Steganotaeniaaraliacea僅具有無規(guī)則型(圖版Ⅰ，G、I)，Eryngiumamethystinum(圖版Ⅰ，E)具3種類型氣孔：無規(guī)則型(30%)，不等型(30%)及平列型(40%)。芹亞科基礎類群僅具無規(guī)則型及不等型(圖版Ⅱ，A～R)。Annesorhizaflagellifolia，A.hirsute及Gliapilulosa(圖版Ⅱ，B～C、K)的氣孔多為無規(guī)則型(75%～100%)，少為不等型(5%～25%)。Bupleurumsmithii(圖版Ⅱ，D)的不等型氣孔(80%)多于無規(guī)則型(20%)。所研究物種的表皮細胞垂周壁通常是直的，但少數物種的細胞壁波狀彎曲，如Astrantiatrifida、Polemanniopsismarlothii及Phlyctidocarpaflava(圖版Ⅰ，D、G；圖版Ⅱ，O)，且同一屬(如Chamarea)的不同物種(如C.esterhuyseniae及C.gracillima)的細胞壁可以是平直或彎曲(圖版Ⅱ，E、F)。

表1　用于研究氣孔結構的傘形科變豆菜亞科及芹亞科基礎類群的憑證標本信息及其采集地Table 1　Taxa of Apiaceae subfamilies Saniculoideae and basal Apioideae examined for stomatal characters, together with voucher specimen details and locality

分子系統(tǒng)學將Polemanniopsis及Steganotaenia兩屬從芹亞科移入變豆菜亞科[6-7,14]，果實微形態(tài)顯示二者果皮中有大量草酸鈣結晶及脊上有大油管，同變豆菜亞科物種很相似[27]，但兩屬依然有芹亞科物種具有的特點，如復傘形花序的特點普遍存在于芹亞科，果實具側脊形成翅存在于芹亞科某些類群(如Heteromorpha)。本研究揭示Polemanniopsis及Steganotaenia具有芹亞科的氣孔特征(表2)。變豆菜亞科的變豆菜族主要有無規(guī)則型及不等型氣孔可以區(qū)別于Steganotaenieae族。刺芹屬的平列型氣孔主要存在于參棕亞科，而牽環(huán)花亞科的氣孔與本研究揭示的氣孔類型相似，主要為無規(guī)則型及不等型。其他刺芹屬物種的氣孔類型需要進一步研究，從而探討此屬同變豆菜族其他類群的關系。

芹亞科植物通常為復葉，但柴胡屬具單葉[28-30]，單葉存在于參棕亞科，牽環(huán)花亞科及變豆菜亞科。此外柴胡屬花粉特征也同這些類群相似，為近菱形，近圓形及橢圓形[31-32]。分子系統(tǒng)研究證明柴胡屬是單系類群，比非洲特有屬進化，但仍位于芹亞科的基礎位置[5,7,33]。果實微形態(tài)顯示所研究的來自非洲的芹亞科基礎類群(Annesorhizaclade，Heteromorpheae，Lichtensteiniaclade，Phlyctidocarpa)的果皮均分布草酸鈣結晶，根據Burtt[34]的描述，芹亞科物種通常沒有結晶，如果有結晶僅存在果實的結合面。本研究顯示除柴胡屬外芹亞科基礎類群的氣孔主要或僅為無規(guī)則型。而柴胡屬屬內物種氣孔結構的變化有待于進一步探究。Choritaenia原位于天胡荽亞科[9]，果實具特殊的油囊[35]，以及氣孔結構均顯示此屬與芹亞科物種相似，支持分子系統(tǒng)學將其移入芹亞科。

表2　傘形科變豆菜亞科及芹亞科基礎類群葉片(苞片或果實)表皮氣孔類型Table 2　Summary of stomatal types of leaf (bract or fruit) epidermis in Apiaceae subfamilies Saniculoideae and basal Apioideae taxa

注：*苞片，**果實

Note: *bract，**fruit

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圖版Ⅰ　傘形科變豆菜亞科植物葉(苞片或果實)表皮氣孔結構(標尺=20 μm)ano.無規(guī)則型氣孔; ani.不等型氣孔; pa.平列型氣孔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

圖版Ⅱ　傘形科芹亞科基礎類群葉(或果實)表皮氣孔結構(標尺=20 μm)ano.無規(guī)則型氣孔；ani.不等型氣孔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

(編輯: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

國家自然科學基金(31270235,31070169)

郭詩琦(1994-)，女，學士，主要從事植物學研究。E-mail:guosq17@163.com

劉玫，教授，博士導師，主要從事植物學科研及教學工作。E-mail:m.r.liu@126.com

Q246; Q944.53

A