王超群，郭詩琦，劉玫，茹劍，王靖茹，程薪宇

(1.哈爾濱師范大學(xué)生命科學(xué)與技術(shù)學(xué)院，黑龍江省普通高等學(xué)校植物生物學(xué)重點實驗室，黑龍江 哈爾濱 150025；2.黑龍江省科學(xué)院大慶分院，黑龍江 大慶 163319)

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傘形科參棕亞科及牽環(huán)花亞科氣孔的結(jié)構(gòu)及其分類學(xué)價值

王超群1**，郭詩琦1**，劉玫1*，茹劍1，王靖茹1，程薪宇2

(1.哈爾濱師范大學(xué)生命科學(xué)與技術(shù)學(xué)院，黑龍江省普通高等學(xué)校植物生物學(xué)重點實驗室，黑龍江 哈爾濱 150025；2.黑龍江省科學(xué)院大慶分院，黑龍江 大慶 163319)

本文首次深入研究了傘形科參棕亞科和牽環(huán)花亞科及相關(guān)類群34屬(參棕亞科9屬，牽環(huán)花亞科21屬)48種植物葉片(莖或果實)的氣孔結(jié)構(gòu)，結(jié)果顯示有3種類型氣孔：無規(guī)則型，不等型及平列型。參棕亞科的氣孔多為平列型，少為無規(guī)則型，而牽環(huán)花亞科多為無規(guī)則型，少為不等型(Gymnophyton除外)。Bowlesia分支的細胞壁多為深波狀，其他分支的細胞壁多平直。氣孔結(jié)構(gòu)的研究為分子系統(tǒng)學(xué)建立的參棕亞科和牽環(huán)花亞科，以及Bowlesia分支提供了形態(tài)學(xué)依據(jù)，并支持分子系統(tǒng)學(xué)將Apiopetalum，Mackinlaya及Stilbocarpa從五加科分別移入?yún)⒆貋喛坪蜖凯h(huán)花亞科，將Homalosciadium，Platysace，Hermas及Klotzschia移出兩亞科。

傘形科；葉；氣孔；系統(tǒng)學(xué)；分類學(xué)

傘形科(Apiaceae)包括天胡荽亞科(Hydrocotyloideae)，變豆菜亞科(Saniculoideae)及芹亞科(Apioideae)[1-2]。Drude[1]主要根據(jù)果實結(jié)構(gòu)將天胡荽亞科(果實具木質(zhì)內(nèi)果皮，沒有心皮柄，油管無或只存在于主脊中)分為兩個族,Hydrocotyleae(分果兩側(cè)壓扁)及Mulineae(分果背腹壓扁)。天胡荽亞科中國僅有3屬(Centella，Dickinsia及Hydrocotyle)[3]，其他類群產(chǎn)于澳大利亞，阿根廷及巴西等地[2]。20世紀末分子系統(tǒng)學(xué)研究表明傘形科，特別是天胡荽亞科，不是一個自然的類群[4-11]。Plunkett[8]指出天胡荽亞科是多元的，原位于Hydrocotyleae族的Hydrocotyle和Trachymene應(yīng)放在五加科(Araliaceae)，Homalosciadium和Platysace構(gòu)成一個分支，為其他傘形科的姊妹群，而其余的天胡荽亞科類群分別位于兩個新亞科，參棕亞科(Mackinlaloideae)和牽環(huán)花亞科(Azorelloideae)。新蘇格蘭特有屬Apiopetalum及生長在所羅門群島的Mackinlaya屬應(yīng)從五加科移入?yún)⒆貋喛?。近期的分子系統(tǒng)學(xué)研究[12]進一步表明參棕亞科(其物種多位于Hydrocotyleae族)包括9屬，分為2個分支：Centellaclade(Centella，Micropleura，Pentapeltis，Schoenolaena)和Xanthosiaclade(Apiopetalum，Actinotus，Chlaenosciadium，Mackinlaya，Xanthosia)；牽環(huán)花亞科(其物種多位于Mulineae族)包括21屬，分為4個分支：Azorellaclade(Azorella，Laretia，Mulinum，Dickinsia，Diplaspis，Huanaca，Schizeilema，Spananthe，Stilbocarpa)，Asterisciumclade(Asteriscium，Domeykoa，Eremocharis，Gymnophyton，Oschatzia，Pozoa)，Bowlesiaclade(Bolax，Bowlesia，Dichosciadium，Drusa，Homalocarpus)和Diposisclade(Diposis)。牽環(huán)花亞科中的Dickinsia屬原位于Hydrocotyleae族，而Stilbocarpa屬原位于五加科。原位于Mulineae族的Klotzschia屬是牽環(huán)花亞科的姊妹群，而另外兩個屬Hermas及Choritaenia靠近芹亞科[12]。

盡管大量研究證明果實形態(tài)特征(包括翅的形狀，分泌結(jié)構(gòu)，內(nèi)果皮解剖特征及結(jié)晶的存在和分布)對傘形科分類是十分重要的[13-16]，可以用于界定參棕亞科及牽環(huán)花亞科[17-18]，并區(qū)分亞科內(nèi)不同的分支[19]，但也有相當多的研究表明傘形科氣孔的結(jié)構(gòu)同樣具有重要的分類學(xué)價值。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(diacytic)及平列型(paracytic)[20-24]。然而迄今未見關(guān)于參棕亞科及牽環(huán)花亞科物種氣孔結(jié)構(gòu)的詳細報道。本研究旨在深入研究參棕亞科和牽環(huán)花亞科所有屬及其相關(guān)類群(Hermas，Homalosciadium，Klotzschia，Platysace)的氣孔結(jié)構(gòu)，弄清氣孔結(jié)構(gòu)是否可以為分子系統(tǒng)學(xué)建立的兩個亞科及分支提供形態(tài)學(xué)依據(jù)。

1　材料與方法

1.1材料

本文所用的研究材料為參棕亞科和牽環(huán)花亞科[11-12]及相關(guān)類群(Hermas，Homalosciadium，Klotzschia，Platysace)，共34屬48種植物的葉片，個別材料因未取到葉片而用莖或果實。材料分別取自澳大利亞國家標本館(CANB-Australian National Herbarium)，澳大利亞國家植物園標本館(CBG-Herbarium，Australian National Botanical Garden)，美國哈佛大學(xué)標本館(GH-Harverd University Herbaria，America)，中國科學(xué)院華南植物園標本館(IBSC-South China Botanical Garden Herbarium)，南非約堡大學(xué)植物及植物生物技術(shù)系標本館(JRAU-Herbarium，Department of Botany and Plant Biotechnology，University of Johannesburg，South Africa)，英國皇家植物園標本館(K-Herbarium，Royal Botanical Garden，England)，美國密蘇里植物園標本館(MO-Herbarium，Missouri Botanical Garden，America)，美國紐約植物園標本館(NY-New York Botanical Garden Herbarium，America)，中國科學(xué)院植物研究所標本館(PE-Herbarium，Institute of Botany，CAS)，西澳大利亞保護和土地管理部植物標本館(PERTH-Western Australian Herbarium，Department of Conservation and Land Management)，摩洛哥科學(xué)研究所標本館(RAB-Herbarium，Institute Scientific，Moroco)及荷蘭烏特勒州立大學(xué)系統(tǒng)植物研究所標本館(U-Herbarium，Institute of Systematic Botany，State University of Utrecht)。研究的材料中僅Centellaasiatica(積雪草)及Dickinsiahydrocotyloides(馬蹄芹)產(chǎn)自中國。物種名稱及憑證標本信息見表1。

1.2形態(tài)學(xué)方法

所有材料浸泡在約90 ℃熱水中約15～30 min，待材料充分吸水膨脹后將材料取出，根據(jù)Zarinkamar[25]的觀察(傘形科植物葉片的氣孔密度，下表皮大于上表皮)，撕取植物葉片的下表皮或莖及果實的表皮，將其放在載玻片上，并在放蓋玻片之前加1～2滴50%甘油水溶液，用具成像系統(tǒng)(Olympus DP 70)的顯微鏡(Olympus BX 51)觀察氣孔結(jié)構(gòu)并照相。每個樣品觀察100個氣孔并統(tǒng)計不同類型氣孔數(shù)量，根據(jù)照片用Photoshop CS 2.0軟件繪制氣孔圖。

表1　用于研究氣孔結(jié)構(gòu)的傘形科參棕亞科和牽環(huán)花亞科植物的憑證標本信息及采集地

2　結(jié)果與討論

根據(jù)前人[20-23,26]對氣孔類型的描述，我們確認所研究的物種均有3種類型氣孔，無規(guī)則型，不等型及平列型(圖1，2)。每個物種具有的氣孔類型及所占的比例見表2。在研究的物種中，部分物種僅具一種類型氣孔(占觀察氣孔數(shù)目的100%)，為平列型(如Actinotusminor，Centellaasiatica及Chlaenosciadiumgardneri，圖1A，C，D)或無規(guī)則型(如Bowlesiaflabilis，Dickinsiahydrocotyloides及Drusaoppositifolia，圖2G，I，M)。一些物種具兩種類型氣孔：多數(shù)無規(guī)則型(80%～90%)，少數(shù)為不等型(10%～20%)(如Asterisciumchilense，Azorellacompacta及Oschatziasaxifraga，圖2A，C，S)；多數(shù)無規(guī)則型(70%)，少數(shù)為平列型(30%)(Hermasquercifolia，圖2Q)；或多數(shù)為平列型(70%～90%)，少數(shù)為無規(guī)則型(10%～30%)(如Apiopetalumglabratum及Micropleuraflabellifolia，圖1B，I)。

早期的研究表明傘形科植物的氣孔多為無規(guī)則型及不等型，少見平列型及橫列型[21-23,26]。所研究的參棕亞科物種僅有或多為平列型氣孔，無規(guī)則型氣孔的數(shù)量較少。在傳統(tǒng)的五加科分類系統(tǒng)[27-31]中，因Apiopetalum及Mackinlaya兩屬的花瓣具爪被放在Mackinlayeae族。Eyde和Tseng[32]及Rodríguez[33]認為Apiopetalum和Mackinlaya兼具五加科(木本及肉質(zhì)果實)及傘形科(花瓣具爪)的特征，Mackinlaya還具有傘形科特有的兩心皮果實及鞘狀葉柄基部，二屬可能為介于兩科之間的類群。果實形態(tài)學(xué)研究[18]顯示Apiopetalum和Mackinlaya的腹束結(jié)構(gòu)同傘形科的參棕亞科物種的腹束結(jié)構(gòu)十分相似，支持分子系統(tǒng)學(xué)[4-5,9,12,34]將二屬從五加科移入傘形科參棕亞科。我們的研究表明Apiopetalum和Mackinlaya(圖1B，F(xiàn)，G)同參棕亞科其他物種(圖1A，C，D，H-L)一樣，主要為平列型氣孔，進一步為二屬應(yīng)位于參棕亞科提供了形態(tài)學(xué)依據(jù)。

表2　傘形科參棕亞科及牽環(huán)花亞科植物葉(莖或果實)表皮氣孔類型

*因未取到葉片而選用莖或果實為材料?！?”表示沒有，“+”表示有。

*The stems and fruits were used in the study because the leaves are not obtained. “-”is absent.“+”is present.

花及果實形態(tài)學(xué)研究顯示原位于五加科，生長在澳洲麥加里島的Stilbocarpa屬的形態(tài)與傘形科物種較為相似[32,35-36]。Mitchell等[37]指出Stilbocarpa同天胡荽亞科的Schizeilema及Azorella兩屬親緣關(guān)系較近，建議將其移入天胡荽亞科。分子系統(tǒng)學(xué)研究[12]將Stilbocarpa置于牽環(huán)花亞科。本研究顯示Stilbocarpa的氣孔主要為無規(guī)則型(圖2BB)，同牽環(huán)花亞科其他物種(如Dichosciadiumranunculaceum及Dickinsiahydrocotyloides，圖2H，I)的氣孔結(jié)構(gòu)極為相似，支持Stilbocarpa屬分類地位的改變。Pozoa，Eremocharis，Domeykoa，Asteriscium及Gymnophyton屬被認為是彼此親緣關(guān)系密切的類群[38]。Henwood等[19]指出這些屬的染色體基數(shù)為x=5。這種較近親緣關(guān)系得到了分子系統(tǒng)學(xué)的證實[12]。本研究顯示無規(guī)則型氣孔是這些屬的主要類型(Gymnophyton除外，需要進一步研究此屬葉片的氣孔)。許多形態(tài)學(xué)研究[17,19,39-40]揭示Bowlesia，Dichosciadium，Homalocarpus，Drusa及Bolax屬均具星狀毛，分子系統(tǒng)學(xué)研究表明他們構(gòu)成一個分支[11-12]。我們注意到這些屬多具有深波狀細胞壁(Dichosciadium及Bolax除外)。這種深波狀細胞壁還存在于Gymnophytonisatidicarpum的果皮，但其他研究的物種的細胞壁平直。

圖1　傘形科參棕亞科及其相關(guān)類群植物葉表皮氣孔結(jié)構(gòu)Fig.1　The leaf epidermis of Apiaceae subfamily Mackinlayoideae and related genera showing the stomatal types　縮寫 Abbreviations: ano (anomocytic)-無規(guī)則型氣孔, pa (paracytic)-平列型氣孔. (A) Actinotus minor, (B) Apiopetalum glabratum, (C) 積雪草Centella asiatica, (D) Chlaenosciadium gardneri, (E) Homalosciadium homalocarpum, (F) Mackinlaya celebica, (G) Mackinlaya macrosciadea, (H) Micropleura renifolia, (I) Pentapeltis peltigera, (J) Platysace maxwelli, (K) Schoenolaena juncea, (L) Xanthosia atkinsoniana. 標尺 Scale bar=20 μm.

圖2　傘形科牽環(huán)花亞科及其相關(guān)類群植物葉(莖或果實)表皮氣孔結(jié)構(gòu)Fig.2　The leaf (stem or fruit) epidermis of Apiaceae subfamily Azorelloideae and related taxa showing the stomatal types 　示無規(guī)則型，不等型及平列型 Including anomocytic (ano), anisocytic (ani), and paracytic (pa). (A) Asteriscium chilense, (B) A. closii, (C) Azorella compacta, (D) A. crenata, (E) A. sessiliflora, (F1,2) Bolax gummifera (1 from middle of leaf balde and 2 from leaf margin), (G) Bowlesia flabilis, (H) Dichosciadium ranunculaceum, (I) 馬蹄芹Dickinsia hydrocotyloides, (J) Diplaspis nivis, (K) Diposis saniculifolia, (L) Domeykoa amplexicaulis, (M) Drusa oppositifolia, (N) Eremocharis longiramea.標尺 Scale bar=20 μm.

續(xù)圖2　傘形科牽環(huán)花亞科及其相關(guān)類群植物葉(莖或果實)表皮氣孔結(jié)構(gòu)Continued Fig.2　The leaf (stem or fruit) epidermis of Apiaceae subfamily Azorelloideae and related taxa showing the stomatal types 　示無規(guī)則型，不等型及平列型Including anisocytic (ani), anomocytic (ano), and paracytic (pa). (O) Gymnophyton isatidicarpum (from fruits), (P) G. polycephalum (from stems), (Q) Hermas quercifolia, (R) Homalocarpus dissectus, (S) Huanaca andina, (T) Klotzschia rhizophylla, (U) Laretia acaulis, (V) Mulinum microphyllum, (W) M. spinosum, (X) Oschatzia cuneifolia, (Y) Pozoa coriacea, (Z) Schizeilema fragoseum, (AA) Spananthe paniculata, (BB) Stilbocarpa polaris. 標尺 Scale bar=20 μm.

Magee等[41]指出產(chǎn)于非洲的Hermas屬應(yīng)位于牽環(huán)花亞科，但分子系統(tǒng)學(xué)研究表明此屬應(yīng)靠近芹亞科[12]，Hermas的葉片具單列多細胞毛狀體，另一產(chǎn)于巴西的Klotzschia(牽環(huán)花亞科的姊妹群)具有長柄的星狀毛，這兩屬植物的毛狀體均與牽環(huán)花亞科植物的不同(未發(fā)表)。Hermas及Klotzschia多具無規(guī)則型氣孔(圖2Q，T)，Hermas具有的平列型氣孔僅存在于參棕亞科。無規(guī)則型氣孔類型普遍存在于變豆菜亞科(Saniculoideae)(如Actinolema及Sanicula)及芹亞科[22-23,26]，而平列型氣孔在芹亞科很少見[22]。筆者認為Hermas的分類地位需要進一步研究。此外本研究還顯示Homalosciadium及Platysace兩屬的氣孔類型(圖1E，J)完全不同于參棕亞科，支持分子系統(tǒng)學(xué)將二者移出胡荽亞科[12]。

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Stomatal structures of the Apiaceae subfamilies Mackinlayoideae and Azorelloideae and their taxonomic value

WANG Chao-Qun1**, GUO Shi-Qi1**, LIU Mei1*, RU Jian1, WANG Jing-Ru1, CHENG Xin-Yu2

1.College of Life Science and Technology, Harbin Normal University, Key Laboratory of Plant Biology, College of Heilongjiang Province, Harbin 150025, China; 2.Daqing Branch of Heilongjiang Academy of Science, Daqing 163319, China

The stomatal structures of the leaves (stems or fruits) of 48 species representing 34 genera of the Apiaceae subfamilies Mackinlayoideae (9 genera) and Azorelloideae (21 genera) and their relative taxa were studied in detail for the first time. Three types of stomata - anomocytic, anisocytic and paracytic - were observed. In Mackinlayoideae, the stomata are mostly paracytic and the minority are anomocytic. In Azorelloideae, stomata are mostly anomocytic and the minority are anisocytic. Cell walls in theBowlesiaclade are deeply waved while they are usually straight in the other clades of Azorelloideae. This stomatal study provides morphological evidence for the two subfamilies and theBowlesiaclade that have been established by molecular research. It also provides evidence to support movingApiopetalum,MackinlayaandStilbocarpafrom Araliaceae to Mackinlayoideae and Azorelloideae, and against placingHomalosciadium,Platysace,HermasandKlotzschiain either of these subfamilies.

Apiaceae; leaf; stoma; phylogeny; taxonomy

10.11686/cyxb2015557

2015-12-09；改回日期：2016-03-25

國家自然科學(xué)基金(31270235和31070169)資助。

王超群(1994-)，女，甘肅張掖人，本科。E-mail:wangchaoqun_723@163.com。郭詩琦(1994-)，女，山東沂水人，本科。E-mail:guosq17@163.com。**共同第一作者These authors contributed equally to this work.

Corresponding author. E-mail:m.r.liu@126.com

http://cyxb.lzu.edu.cn

王超群, 郭詩琦, 劉玫, 茹劍, 王靖茹, 程薪宇. 傘形科參棕亞科及牽環(huán)花亞科氣孔的結(jié)構(gòu)及其分類學(xué)價值. 草業(yè)學(xué)報, 2016, 25(10): 132-141.

WANG Chao-Qun, GUO Shi-Qi, LIU Mei, RU Jian, WANG Jing-Ru, CHENG Xin-Yu. Stomatal structures of the Apiaceae subfamilies Mackinlayoideae and Azorelloideae and their taxonomic value. Acta Prataculturae Sinica, 2016, 25(10): 132-141.