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(2) Y.. 36. the. paper,. are. tions. may. have. intended. scanning. to. There. are. followings. In the. intraspecific. many. (2) H.. (3). H.. (4). H.. (5). H.. 1988.. macroPhylla. var･. macroPhylla. var･. acuminata. macroPhylla. var･. angusta. the. on. meager. ornamentally. islet,. Enosbima in Mt.. for minata of T6hoku. in 1987,. These. were. collected, fully-grown leaves･ their of. City,. and. from. the. blades.. trichomes leaves. The. individuals. collected from Prefecture,. from. ones. were. leaves. of them. were. Wild. forma. acu-. Laboratory Yamanashi. Kiyosato, grown. and. planted. The. at. planted. Haccbou. near. wild. and of forma megacarPa Mt･ Hakkoda Botanical at and. macro-. growlng. ones. growing. very. were. forma. of. from. on. the. campus. from. collected. square millimeters about丘ve in 2･5% glutaraldehyde vere丘Ⅹed pieces of leaves After buffer 7･217A)･ (pH phosphate each in O･1M. into. sectioned. dehydrated. was. material. because. leaves. foliage. young. cuttings･. collected The small. tetroxide,. very. in. November. to. September. foliage. Cauline. shrub･. were. Twigs. were. osmium. the丘Ⅹed. and. investigated･. macroPhylla. soft-wooded. collectedノfrom forma normalis･. They respectively･ Universlty･ National. leaves. razor. 1%. were. macro♪hylla･. was. angusta. deciduous,. respectively.. grown. a. using. normalis. were collected cuttings Prefecture in 1983 Aomori University,. of the Yokohama the individuals. and. a. making. prefecture. group. acuminata. forma. leaves. Fujisawa. Amagi,. We. megacarPa. forma. angusta and forma normalis City, Kanagawa in Fujisawa. phylla, forma. with. surface. forma. megacarpa. is. forma. forma. macrophylla. fully-grown. their. varia-. group･. in Hydrangea. of their leaves. macroPhylla. var･. macroPhylla. as. well. intraspeci丘c. some. macroPhylla. entities. structure. var･. macroPhylla. bad fully grovn vhich But, in forma macroPhylla. leaves. botanical. macroPhylla. Hydrangea. that presented the intraspecific. macroPhylla. H.. transverse. the. arLd班ethod. the surface. of them,. (1) Hydrangea. among. H･. of. surface. and. group,. that. of. in. (SEM)･. microscope. electron. leaf. inferred leaves. of. view. the. observe. has. he. nlaterial. p.nd. found. described. Furthermore, entities. in the surface occur. botanical. at. TANAKA. leaves. section. as. M.. and. characters anatomical of foliage of Hydrangea macroPhylla even differences in foliar distinct anatomy. present. there. has. SAT6. in. a. graded. ethanol. series,. ethanol. was. re-. was critical-point dried using carbon material a on fluid. The leaf pieces were stub using as mounted dioxide the transition The coated material a tape, with platinum･ double-stick and sputter-coated electron microscanning was with a JEOL JSM T-loo viewed and photographed. placed. with. scope.. The. were. length from. measured. Cosmozone. lS).. The. acetate･. amil. and. the. height. photographs. of using. structures. the. image. and. the. area. analysis. of cell surface program. (Nikon.
(3) Leaf. Hydrangea. of. surface. 37. macro♪hylla. Observation. (1) Stomata Stomata leaves. (Figs. 1A,. of丘ve. forma. members. ones. on. September.. the. There. was. There. outer. ledge Table. 1.. 4A). macroIIhylla. forma. normalis. were. the. on. absent. immature. the. on. only. adaxial. (Fig. 2B). stomata. at the of the leaves collected in difference structure of the stomata. no. however,. (〃m). length. of. in. in Table. shown. as. aperature. difference. a. outer. length. l･. The. ledge. stomatal. 10. 7. 6.2. 16.8. 9. 9. 5.7. 15.2. 6.9. 16.6. acuminata. 16. 2. 10.4. 22.9. forma. angusta. 13. 9. 8.1. 19.8. var.. forma macro?hylla Stoma. B. Hydathodes. macro♪hylla C,. normalis D). ,(B, C. Adaxialepider申41S.. A.. D.. margin. tively. irr喝ular. trichom色こ--(s) with. epidermis.. abaxial the明年rly. smoqth. macroPhylla on. Nearly. abaxial. F.. of. smooth Trichomes. surfaceヤ. very. any. of their of. length. forma. Hydrangea. Of. length. Maximum. megacarpa. 1.. the. aperture･. forma. forma. In. among. ll.9. Fig.. side.. beginnlng. average. length. Minimum. of. among. length. long. the. side. abaxial. surface. was,. Long. present. they. few. a. Average. forma. were. but. were. abaxial. 丘ve members. stomatal. 3A,. examined. there. megacarpa,. mature. 2A,. tooth. (A,. E,. surface. epidermis･ young. leaf,. F). and. of leaf E･ Rela-. Npte. a.
(4) 38. Y.. SAT6. M.. and. TANAKA.. forma is similar to that in forma Their length average macrophylla normalis･ is relatively it is abotlt two-thirds short the length and of forma acuminata has longest aperture which the The among all members measured･ average length. of. length. among. The few. leaf. the. leaves. that. ゝnOt. The. elevated. but. the. The. veins. In face. view. the In. looked. leaves. were. three. abaxial. average. adaxial. relatively angusta. and. are. about. both. f6rma. to. very. and. the. and. have. examined. a. a. one. forma. relatively had. meshes the. veins.. normalis,. the. epidermal. relatively. of. with fdrma. enclosed. areoles. slightly. face. the. areoles.. associated. associated than. view. the. with. shown. are. cells. and. in. outline of Table 2; the. and forma megacarカa are forma forma acuminata and. normalis. while. those. another,. too･. The. former.. average. The. (Figs. 1C, D). several. cells. in. outline. area. in. normalis, irre即1ar. epidermal. simpler. values. of the. values. pinnate, delimltlng Small. exposed. whole. latter. surface. of. in rather smooth former, some of the. was. usually. the. thoughin. of the. values. In the. (Fig. 1E).. abaxial epidermis folds at their exposed. (Fig. 1A).. surface leaves. adaxial and abaxial epidermis of of forma forma (Fig. 4D), the outline of each epidermal angllSta was or areoles lobed. Each complicate and deeply slightly. vith. of thei,r adaxial (Figs. 3B, 4B) had many epidermaトcells its exposed In forma and ridges on surface. acumiあata,. (Fig･ 3B), while Tabie. 2.. †informa. Ar甲(FLm2) epidermal. a?gusta,. enclosed cell. with. associated. Average. area. forma. macrophylla. 884. 0. forma. normalis. 710.7. forma. the. (Fig･ 3D) and. cell associated. continuous. the. along. wa､s. the. other, to. examined. along. associated. subsidiary cells usually In face viev of both. acuminata. along. teeth. with. sides,. abaxial epidermis. macrophylla. leaves. enclosed. abaxial. mqcrophylla,. each. similar times. adaxial. of. elongated. The. cell. in forma. two. surface. all. teeth. not.. formed. forma. had.. members. similar are. some. of. crater. or. members. veins. and. narrow. epidermal. values. a. open. all. of branched their. forma. other. each. of. value. In all members,. margln･. surface the. like. be. toothed. a. on. present. might. a. on. p･resent. were. crater. both. veins. they. of and. on. were. macro♪hylla and the areoles (Fig. 1C) had. the. intermediate. the. cells. venation. secondary. had. examined. hydathodes. rim, Epidermal. (3). with. is. angusta. measured.. (Fig･ 1B). leaf margln･. two,. members. of all members. hydathodes margln,. with. all. forma. and. megacarpa. Hydathodes. (2) a. forma. megacarpa. 1527.7. forma. 1498. 3. angusta一. outline with. each. into small of each. or. ridges short. was. ridges. abaxial. Maximum. area. 632. 3. 1192. 1. ､､523. 6. 889. 5 :･”. 407.8 I. of. grooves. areoles.. Minimum ＼. 736.6. forma.acuminata. it broke. fine distinct. 792.0･ 733.4. L. 957.7 2601.7 2219.4. area.
(5) Leaf. Fig.. 2.. Hydrangea. Hydathode.. their. these by. the. grooves. epidermal. fixation,. angusia,. i･nvested. with. smooth･ bad. thin. some. those. at. These. the. surface. folds. seem. drying. both. In the. cells. on. and. they. greatly. surface. of. surface. surface. are. of abaxial. cell甲d. the. (Figs･ 3A,. 4A)･. of these cells in forma to be artificially. lformed. forma. and. acuminata. adaxial. and. swelled. the. solved. they. or. epidermal had not usually. subsidiary. folds at their exposed. or/and. whole. is, the the. of. of the be. abaxial. (Fig. 4C). That. E.. base.. cell val上 itself. angusta. and. epidermis.. trichome. every. B.. grooves. without. abaxial at. hand,. other. Stoma.. surface appearance it remains to. the. surface.. normalis.. the. by. forma. and. to. similar. ce11■ wall,. Therefore,. C).. the. several. all epidermal a. On. Howev?r,. dehydration. almost. formed. ridges. (s). structure. complicate Hovever,. are. its exposed. almost forma. and. more. had.. (Fig. 3C). them. were. macrophylla. abaxial. outwardly on. epidermis. both. sides. forma were. (F､igs. 3B,. sides. vas. lrregular.. In face on. to. bad. vax.. on. were. cells. folds. during. the. epicuticular. Hillock,like. A.. megacar♪a.. cells epidermal Relatively lrregular. D.. former. and. ridges. adjacent. very. than. acuminata. and. latter. forma. Note. F.. surface.. the. grooves. cell of forma. c, 4B,. adaxial. epidermis. whether formed. cells Their. on. 39. macroPhylla. megacar9a. surface.. exposed. Therefore,. (Fig. 4B). adaxial. var. macro♪hylla Adaxial epidermis.. C.. at ridges Trichomes. of Hydrangea. surface. both. adaxial epidermis. sides and. of forma. view was. abaxial was. megacarpa,. deeply. or. similar. to. slightly was. epidermis that. also,. rathet found. the. lobed.. outline of The whole. irregular･ occasionally. The in. the. cells epidermal both the surface･-of. irregularity forrrla. of abaxial ma占Irophylla･ In.
(6) 40. Y.. Fig.. 3.. Hydrangea. Irregular radially. Fig.. 4.. cells. arranged. (*). at. macrophylla. epidermis. cells. C.. epidermis.. Hydrangea. adaxial. var.. macroPhylla. adaxial. arran宮ed. SAT6. C.. (*) at. M.. TANAⅠくA. forma. acuminata. Irregular. abaxial. A.. acuminata.. epidermis.. D.. Stoma･. Tricbome. B.. with. its base.. Far. angusta. Irregular its base,. and. forma. A.. angusta.. abaxial･･epidermis.. D.. Stoma.. Trichome. B. Irregular with. radially.
(7) Leaf. forma and. That. ridges.. of forma. that. however,. of the were. that. forma. of. had. without. absent. crossing. exposed. Thus,. in forma surface of leaf epidermis megacarpa to forma as that of forma well as macrophylla. has. of. forma at. surface it seems. adjacent similar. megacarpa, grooves. which the. that. features. with-. more. was. In. acuminata.. grooves. continuous. megacarpa. (Fig. 2B).. fine. many. was. ridge. the. cells. epidermal. partially. and of forma. ridges. condition. than. angusta. some. ridges. surface. had. cells. epidermal (Fig. 2B). The. short. or. 41. macrophylla. its adaxial. surface. small. is, the. Hydrangea. of. usually. exposed. into. breaking. any. and. their. at. ridges. out. to. also,. megacarpa,. surface. exposed. to. common. that. of. angusta.. (4) Trichomes Trichomes, more. usually,. distributed. were. usually fully-grow.a. leaves. the. mature usually. 150-250fJm. long.. were. face. many. But,. common. the. at. all members. examined.. The. The. trichomes. forma. and. surface.. higher. were. of forma. protuberances. those. of. of forma 4FLm. about. leaf. areas. other had not. as. trichome as. well. had. On. the. 200〃m. trichomes. and. long. acu-. forma inter-. and. the. in. and. hand,. of veins and. sur-. forma. other. long. was. the. at. 300FLm. than. rarely,. ones. a. the surface. on. (Fig･ 1F),. (Fig. 2E). more. than. for. trichomes. meager･. forma. (Fig. 2E),. megacarpa (Fig. 4D) invariably. than. protuberances. were. forma. of. angusta. The. in length. on. COmmOnly. long.. less. they. and. examination. viewed. areas. megacarpa. trichomes. meager,. angusta,. most. usually. were. usually. 200fLm. than. more. with. trichomes. and interveinal. 800〃m. forma. tubercular. the. forma. were. to. not. among. In forma. In. our. developing. of the forma megacarPa. surface. (Fig. 3D), their trichomes minata the former, some of them attained had short angusta (Fig. 4D) usually veins.. protuberances. acuminata. surface.. any examined A trichome surface. of their it was at its surface and. leaf. trichomes. were. leaves. the. young. they. very. In. were. of. in their. In particular,. veins.. were. of vein.. the. of. small. megacarPa,. of the. among. some. smooth-walled forma. nearly. relatively. trichomes. area. whole. had. In forma. the. tw`o,. the. leaf usually. there. surface. leaf and. adaxial. they. the. Furthermore,. through. trichomes. only. in. leaf surface.. of the. on. the. on. abaxial. forma. and. macrophylla. of the. surfaces. than surface however, normalis,. the. on. prevalent. both. on. occurred. had. many. small. angusta;. usually,. highand. latter. the. at. protuberances. forma. and. megacarpa. (Fig. 3D). acuminata. the. former. had. those. acuminata. usually had the. usually. of less. their. than. 2FLm. high. There base was. of. was. the. base. appearlng angusta. of the base.. trichome In. In. trichomes.. surrounded level. difference. a. with. forma. several. (Fig. 4D),. cells the. epidermal these. hillock-like. a. megacarpa,. epidermal. appearance. megacarPa. And. epidermis. That is,. forma. in the. at. trichomes their base.. epidermal. cells. of epidermal (Figs. 2E, F),. cells. the. to. turgid. were. which. situated at the base of trichome. cells. raise. above. turgid. the cells arranged radially around was formed at the trichome structure on. the. In forma at. the. vein. also. had. a. ring. of turgid-. (Fig. 3D) and acuminata base of their trichomes. forma in the.
(8) Y.. 42. interveinal That. areas. is, they. no. form. to. radially hillock-like. TANAKA. a. base. of trichomes. were. from. the. the. members. are. present. the. trichome absent. were. structure. and. turgid･. not. In this respect,. different. were. in. vein. they. base･. their. at. structure. leaf. the. on. but. rlng,. angusta and forma acuminata Special cellular arrangement megacarpa,. of forma the. bad. M.. and. of forma. trichomes. at. arranged. SAT6. for. except. forma. megacarPa.. Disc11SSion ln. all. are a. few. the. on. present. features be. can. is. there. and. epidermis. here,. examined. members. difference. no. surface. teeth. of. have. to. H.. A:. B:. members leaf. fully-grown. macrophylla. acuminata. ∀ar.. angusta. the. ′rhe whole lS. usually gular. trichomes. nearly. epidermis hand,. three. in. not. any. epidermis. in. surface,. though. the. continuous. ones. adaxial. of forma. cells. whole form.a. and. epidermis have. B. features attention to. A. the. has. three. has. an. of the. two. that. format. trichome. common. members. cell. of. abaxial. and. features of Group. and That. of A.. assigned. to. Every. lS,. of. Group. B. has. the. surface･. acuminata also. the. of Group. three. the. exposed. from. their. forma. members. surface which How`ever, it should. the. epidermal. megacarpa the. respects,. some. at. ridges. leaf. other They of. the. differ. But,. three. the. cell. on. acuminata. epidermis in forma. adaxial. On common･. ridges. angusta. and forma have･not grooves and. irre-. rather. surface･. and. tool. members. their. of. relatively. forma. two. with. surfaces･. grooves. irregular,. megaCarPa. are. their. at. surface･ In these epidermis.. abaxial some. trichomes. irregular. rather. have. ridges. ridges. megacarpa. is distinctly. Every. surfaces,. present,. the. leaves. its exposed. at. their. rarely of. are. occasionally. many丘ne. partially both of adaxial. angusta. epidermis. B,. discontinuous. surface. abaxial. protubelances. megacarPa. of Group. those. small. of forma. irregular. an. Group. and. are. of their. surface. on. protubelances. macrophylla.. of Group. many. small. the. on. surface. and. forma. grooves. members. have. invariably. The. in. angusta. meager. smooth. thoughthere. smooth,. follows:. as. groups. acuminata. forma. many. which is found. megacar♪a. forma. very. not. examined. two. forma. are. nearly. adaxial. ab耳Ⅹial epidermis has. have. a. with. of the. surface. trichomes. are. signi丘cance. normalis. megacarpa. usually. diversity. macro♪hylla. var.. They. the. macroPhylla. macrophylla macrophylla A, of Group. there. forma. var.. H.. Thus,. macrophylla. macrophylla. abaxial. Hydathodes. forma. ∀ar.. H.. In two. of all members divided into. are. macroPhylla. H.. and. phylogenetic. ∀ar.. H. Group. and. the. at. type.. However,. examined.. of leaf. Group. though. along. structure surface kinds examined of plants. Five. structure. only. the leaf margln.. taxonomic. a. the. among. in their. to all members. common. expected. stomata. B of. members. differ from received features. the much. common. A.. with. a. hillock-like. structure. is prevalent. throughsection. Hy-.
(9) Leaf. drangea. of the. is present. only. forma. and. Bl. Subgroup. B2:. The. rare. which some. of the. surface closed. length.. macroi)hylla. var.. angusta. in section. Hydrangea. a. epidermlS, of the. features B. in Group. from. that. macrophylla. and. three. other. there. of shows found in leaves of all members features to show seems that taxonomic are. or. ones. names. be. of. other. and. species. features detail. the. of which. the. 広義のアジサイ(Hydrangea. feature. a. detail. the. and. examinafeatures. have. structures. in. used. We. our. that. examination. and. comparison. of. these com-. to. assigned. various. a. paper. consider. entities. in. illtermediate. some. names. intraspeci丘c. Hydrangea. 摘. in its. different. precisely. of foliar. in the. (1965).. on. examination genus. Occurrence. scienti丘c. Ohwi. the. among. are. normalis. The. based. areoles. value. (Sat6, 1989). This examined intermediate features among. by. en-. A.. forma. slgni丘cance.. exposed. the. link. to. seem. in. areas. of. with intermediate. an. to. megacarpa, their. at. values. small. megacarpa. here. examined features found the. chosen in future,. In forma. ridges. seems. Corm'dia. cells associated. has. in Group. some. been. reviewed. many. macroPhylla among. have. which must. parison. phylogenetic. the. Of. aperture. members. are. in section. (Stern, 1978). and. A. in Group. that. with. in forma. with. that. ours. found. angusta.. surface. epidermal. ledge. megacar♪a acuminata. forma. in splte. follows:. as. forma. the grooves. not. and. even. forma. smooth. surface. abaxial. stomatal. outer. forma. of. with. smooth have partially. These. anatomy. tion. H.. outline. of leaf surface foliar. macrophylla. aFuminata. has. the. view,. Both. H･. adaxial. the. with. in face. megacarpa. of trichome. of the trichomes in forma acuminata to forma megacarpa.. subgroups. var.. cells. epidermal. two. ∀ar.. feature. trichome. a. this type. common. macrophylla. occurrence. trichome. a. features. salient which into B is subdivided. H.. :. But,. it is absent. and. megacar♪a,. 43. macrophylla. (Stern, 1978).. have. Group. Subgroup. long. forma. angusta. In this respect,. link. Hydrangea. genus in. of Hydrangea. surface. 〟.. features. is composed.. 要. macroilhylla)には多くの種内分額群が記載されている｡その内,. 次の5種類の成熟した菓の表面構造について比較することを試みた｡ (1)アジサイ. H.. macro少hylla. (2)ガクアジサイ. H.. (3). H.. -ゾアジサイ. (4)ヤマアジサイ (5)アマギアマチャ. H.. var.. macro♪hylla. macroPhylla var.. macrophylla. ∀ar.. macrophylla. ∀ar.. H.. macrophylla. var.. macrophylla megacarPa. forma. macroPhylla forma. forma. normalis megacarPa. acuminata. forma. acuminata. angusta. forma. angusta. アジサイ(狭義)ほガクアジサイを鑑賞用に改良して作り出されたものと言われているが,両者の 葉の表面の構造には特筆すべき特徴の違いは見いだせなかった｡しかし,他の3種類の菓の表面構 造は,明らかにアジサイ(狭義)やガクアジサイとほ異なった特徴を示していた｡アジサイ(狭義) とガクアジサイの葉にほ毛状体はほとんど見られず,向軸側の表皮細胞の表面はほぼ平滑で特別な 構造ほ見いだせなかった｡しかし,. -ゾアジサイやヤマアジサイ,アマギアマチャの菓にはかなり よく毛状体が見られ,向軸側の表皮細胞の表面にほ特徴的な非常に細かい畝と清からなる縞模様が.
(10) 44. Y.. SAT6. M.. and. TANAKA. 見られたoエゾアジサイの毛状体ほ,今回調査した他の4種類では見られない小丘状の構造をその 基部に持っていた｡この小丘状の構造を持つ毛状体ほ,アジサイ(広義)の収容されているアジサ 1978)｡つまり,小丘状の構造のない毛状体がアジ. イ節にほ広く存在しているものである(STERN,. サイ(広義)でほ普遍的であり,小丘状構造を持つ毛状体は少ないという点では,アジサイ(広義) はアジサイ節のなかで特殊な存在であり,アジサイ(広義)の中では小丘状の構造を持つ毛状体を 持つ-ゾアジサイは特筆されるべき存在である｡この論文では大井(1965)によって採択されてい I7ydrangea. る学名を用いたが,. macro♪hyllaという学名の下にまとめられている分類群の分類学. 的な扱いは,アジサイ属の種間やアジサイ(広義)の種内での植物学上の多くの特徴を調査したの ちに,再検討されるべきであろう｡ この研究にほ,昭和63年度の教育研究特別経費(｢電子顕微鏡を利用した教材開発のための総合 的研究+一研究代表者:松田忠男教授-)から一部援助していただいたことをここに付記し,感謝 の意を表したい｡. Reference良 AIRY. SHAW,. H.K.. WILLIS, A.. CRONQUIST, bia. HooKER,. zenfamilien, ∫.D. 1865. l.. Loovell. &. 18a:. ed. &. &. ferns･. t)y. J･C･. of classi丘cation. of且owering. Colum-. plants･. In A･. ENGLER. &. K･. PRANTL,. Die. naturlichen. Pflan-. 74-226. In G.. BEN′1､11AM. J･D･ HooKER,. &. Genera. plantarum. London.. Company,. The. Press,. families. of. flowering. plants.. Vol.. Ⅰ･. 2nd. Dicotyledons,. ed･. oxford. into. of Japan. (transferred the the Japanese Inst., Smithsonian Washington, D･C･. Flora. WALKER). E.H.. system. plants. London･. York,. New. Saxifragaceae.. J. 1959.. J. 1965.. integrated. flowering. the. Press,. University. Saxifragaceae. 2nd.. Reeve. Clarendon ofIWI,. An Press.. 1928.. A.J.. HuTCIiINSON,. sAT6,. ed.. University. ENGLER,. Cambridge. 1981.. dictionaryof. A. 1973.. (Reviser). 8th. English. by. F･G･. MEYER. Hydrangea. group･ (in JapamacroPhylla : 15-26･ English with summary)横浜国立大学理科教育実習施設研究報告5 HySaxifragaceae, and W.L. 1978. Comparative anatomy systematics of woody. Y.. Anatomical. 1989.. study. of. leaves. of. nese. STERN,. drangea. A.. TAKHTAJAN, Bot. TIIORNE,. Bot.. 46:. Rev., R.F.. Biology,. 1976. 9:. Journ.. 1980.. Soc.,. Linnearl. Outline. of. the. 76:. 831113.. classi丘cation. offlowering. classification. of. plants. (Magnoliophyta)･. 225-359. A. 35-106.. phylogenetic. the. Anglospermae･. Evolutionary.
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