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Variation in Shoot Organization in Prunus (Rosaceae) with Special Reference to Prunus ogawana PDF

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植物研究雑誌 J. Jpn. Bo .t 76: 329-338 (2001) Variation in Shoot Organization in Prunus (Rosaceae) with Special Reference to Prunus ogawana Akitoshi IwA MOTO\ Keiko MIYA ZA阻band Jin MURATAc aUniversity Museum,th e University of Tokyo, Hongo 7-3-1,Bu nkyo-ku,T okyo,1 13・0033JAPAN; bMakino Herbarium,T okyo Metropolitan University, Minamiosawa 1-1,Ha chi吋i,Tokyo,1 92-0397 JAPAN; [Present address: Matsubara 3-39-29,Se tagaya-ku,T okyo,1 56-0043 JAPAN] CBotanical Gardens,Gr aduate School of Science,th e University of Tokyo, Hakusan 3-7-1,B unkyo-ku,T okyo,1 120001JAPAN ・ (Received on May 11,2 001) To determine variation in shoot development in Prunus (Rosaceae),w e observed and compared the shoot organization of 13 species (9 species of subgen. Cerasus sect. Pseudocerasus; 1s pecies of sec .tMicrocerasus; 1s pecies of sect. Phyllomahaleb; 1s pe- cies of subgen. Prunus; and 1s pecies of subgen. Padus). As ar esult,w e can categorize the observed species into four distinct groups based on shoot organization of 1) types of winter bud growth,2) growth of buds in the axils of scale-like prophylls,an d 3) shoot ab- scission. The first group includes all species of subgen. Cerasus sect. Pseudocerasus and sec .tPhyllomahaleb except P. ogawana. The second group includes only P. ogawana,th e third group contains P. mume and P. glandulosa,a nd the fourth group contains only P. grayana. Prunus ogawana (subgen. Cerasus sect. Pseudocerasus) is distinct from other members of sect. Pseudocerasus in that some winter buds develop into mixed shoots pro ・ ducing both flowers and leaves,a nd some buds in the axils of scall-like prophylls grow into vegetative shoots. In addition,t his grouping is basically supported by the phylogenetic DNAa nalysis of Prunus. Key words: Prunus,P runus ogawana,P runus subgenus Cerasus,sh oot organization Introduction to determine the degree of variation between Prunus (Rosaceae),c omprising about 200 them. Wee specially focused on P. ogawana species,ha s been usually classified into sev- Makino (subgen. Cerasus sect. Pseudocera- eral subgenera (Rehder 1940,G ohra and sus). The species usually blossoms within Panigrahi 1995). Ohba (1989) recognized three ye sof germination (Makino 1906), 訂 five subgenera (Prunus,A mygdalus,P adus, while other species of Prunus usually need Laurocerasus,C erasus) in Jap an. There are 10-15 years for flowering. Additionally,th e some references to shoot organization in species produces fertile seeds that can rela- Prunus in the descriptions of the species,bu t tively easily propagate. Hence, Prunus critical comparative studies have not been ogawana was expected to be suitable for ex- conducted. Weo bserved the shoot organiza perimental studies of flower formation in 四 tion of species of 13 Prunus (mainly Prunus using genetic recombinant methoas. Jap anese wild species) to comp訂ethem and Flowering at aj uvenile stage has been -329ー 330 植物研究雑誌 第76巻 第6号 平成13年12月 considered to be the only distinct characteris- subgen. Cerasus sect. Phyllomahaleb (P. tic differentiating it from other species of maximowiczii). The materials used in this sect. Pseudocerasus. Our preliminary obser- study eshown in Table 1. Weo bserved de- 訂 vations revealed that all winter buds develop tails of the shoots macrographically. A into shoots with flowers. This feature does stereoscopic microscope was used to observe not occur in other species of Prunus and axillary bud organization. shows that the shoot organization of P. ogawαna may be unique. Detailed observa- Results tions of shoot organization in P. ogawana Fe atures observed and of other phylogenetically closely related We determined that伽 eefeatures were species of Prunus 'enecessYto ascertain important for analyzing shoot organization; 紅 紅 that the feature is species-specific. (1) type of winter bud growth,(2 ) growth of buds in the axils of scale-like prophylls and Materials and Methods (3) shoot abscission. Observations of each We observed shoots of 13 species of item eas follows: 紅 Prunus throughout two years (from April 1998 to March 2000) using living trees and (1) Types of winter bud growth sample branches obtained from the ees. All the species examined in this study e 佐 訂 Among 13 species,ni ne belong to subgenus deciduous and all new shoots extend from Cerasus sect. Pseudocerasus,th e other four winter buds. The winter buds of species of belong to: subgen. Prunus (P. mume), Prunus usually develop into one of three subgen. Padus (P. grayana),su bgen. Cerasus types: (1) av egetative shoot that produces sect. Microcerasus (P. glandulosα) and some leaves and no flowers,(2 ) ar eproduc- Table 1. Observed materials of Prunus s. 1. in this study. Scientific names e rangedaccording to 訂 紅 Ohba (1989) Subgenus Section Species Source* Cerasus Phyllomahaleb maximowiczii Rupr. IV Pseudocerasus apetala (Siebold & Zucc.) Franch. & Sav. w cerasoides D. Don v訂.campanulata (Maxim.) Koidz. w incisa Thunb. Y w E jamasakura Siebold ex Koidz. AY 11 111 w E - pendula Maxim. f. ascendens (Makino) Ohwi TE w l sargentii Rehder y- w - speciosa (Koidz.) Nakai w verecundiα(Koidz.) Koehne 111 w ogawana Makino 11 Microcerasus glandulosa Thunb. IV Prunus mume (Siebold) Siebold &Z ucc. IV Padus grayana Maxim. IV *So urce. 1: Tokyo Metropolitan University. 11: Botanical Gardens Koishikawa,G raduate School of Science,t he University of Tokyo. 111: Botanical Gardens Nikko,G raduate School of Science,t he University of Tokyo. IV: Tama Forest Science Garden. December 2001 Joumal of Japanese Botany Vo .l76 No. 6 331 F - p reproductive shoot reproductive shoot (a) (b) vegetative shoot vegetative shoot (c) (d) Fig. 1. Types of winter bud growth in Prunus s. 1. (a) Winter buds grow into both reproduc- tive shoots and vegetative shoots. (b) All winter buds grow into shoots with flowers. Some shoots also develop leaves (mixed shoots). (c) Winter buds grow into only vege- tative shoots. (d) All winter buds grow into shoots with leaves. F: flower. B: bract. L: leaf. S: scale-like leaf. P: scale-like prophylls of winter buds. Arrows indicate indefinite shoots. 332 植物研究雑誌第76巻第6号 平成13年12月 tive shoot that produce some flowers and no (c) Growth into only vegetative shoots leaves or (3) am ixed shoot that produce both All winter buds grow into vegetative flowers and leaves. We also found that the shoots in this type and no shoots with flow- species of Prunus produce three kinds of ers were found. Buds in the axils of scale 田 winter buds in various combinations,w hich like prophylls also show shoot elongation of are illustrated in Fig. 1. another type (see (2) Growth of buds in the (a) Growth into reproductive shoots and axil of scale-like prophylls). The species vegetative shoots which show this type are P. mume and P. Winter buds produce ar eproductive shoot glandulosa. and av egetative shoot in this type. No buds (d) Growth into vegetative shoots and develop into am ixed shoot. An apical bud is mixed shoots more likely than al ateral one to grow into a All winter buds develop leaves and some- vegetative shoot. The species that show this times also produce some flowers. The spe- type are P. jamαsakura,P . verecunda,P . cies showing this type is P. grayana. sargentii, P. speciosa, P. pendula, P. cerasoides,P . incisa,P . apelata and P. (2) Growth of buds in the axils of scale-like maxzmowzcZll. prophylls (b) Growth into reproductive shoots and Some species of Prunus have buds in the mixed shoots axils of scale-like prophylls of the winter All winter buds produce flowers. In addi- buds (we call the bud in the axil of al eaf the tion,s ome of them also develop some leaves, main bud to distinguish it from the buds in but buds in the axils of scale-like prophyll the axils of scalelikeprophylls) (Fig. 2). 田 show shoot elongation of another type (see The prophyll buds do not appear to expand (2) Growth of buds in the axil of scale-like soon,b ut as tudy by Guo et a .l(1992) im- prophylls). The species showing this type is plied that such buds in P. persica regully 紅 P.ogawana. develop into shoots. Our observations re- vealed that the axillary buds show av ariety … Aowers s (altemate) Buds in il 鉱 of as cale-like prophyll Fig. 2. Structure of winter bud (a mixed bud) in Prunus s. LF lower buds produce no leaves and vegetative buds produce no flowers. Meaning of symbols are same as those in Fig. 1. December 2001 Journal of Japanese Botany Vol. 76 No. 6 333 (a) r--h、., ‘、u J \判各例舛~ L ‘ (d) Fig. 3. Growth of buds in the axil of scalelikeprophylls in Prunus s. l. (a) Buds in the axile of scale-like 回 prophylls do not develop. (b) Buds in the axile of scale-like prophylls of mixed shoots develop into vegetative shoots. (c) All buds in the axile of scale-like prophylls develop into reproductive shoots. (d) Buds in the axil of scale-like prophylls develop vegetative and mixed shoots in following year. Shoots indicated by broken lines extend one ye aftershoots in solid lines develop. Shoots within solid ellipse 訂 ederived from main buds; shoots in broken ellipse extend from buds in axil of scale-like prophylls of 訂 main buds. Meaning of symbols are same as those in Fig. 1. of growth pattems,fo r which we can distin- shoot. Shoots which have no leaves were guish four types (Fig. 3). more likely to develop these axillary vegeta- (a) Bu ds in the axils of scale-like tive shoots than shoots with leaves. A single prophylls not developing bud in either axil of two scale-like prophylls Axillary buds were recognizable rarely in grows into av egetative shoot. The species the axils of winter scale-like prophylls,or if showing this type is P. ogawana. they were present the bud hardly expands. (c) A ll buds in the axils of scalelike 回 The species exhibiting this type are prophylls develop into reproductive shoots P. jamasakura,P. verecunda,P. sargentii, Almost all buds in the axils of the scale- P. speciosa,P . pendula,P . cerasoides,P. like prophylls grow into reproductive shoots. incisa,P. apelata and P. maximowiczii. Buds in both axils of two scale-like (b) Some buds in the axils of scale-like prophylls expand. They also grow into prophylls develop into vegetative shoots shoots in the same ye whenthe main buds 紅 The bud in the axil of scale-like prophylls developed. The species showing this type are of an ew shoot sometimes grows into av ege- P. mume and P. glandulosα. tative shoot in the same year as the new (d) All buds in the axils of scale-like 334 植物研究雑誌 第76巻 第6号 平成13年12月 prophylls develop into new winter buds while all vegetative shoots persist to develop Nearly all buds in the axils of the scale- new shoots from their axillary buds in the like prophylls develop into winter buds that following year. The species showing this grow into vegetative or mixed shoots in the type are P. jamasαkura, P. verecunda, following year of main bud expansion. A P. sargentii,P . speciosa,P . pendula,P . single bud in either axil of two scale-like cerasoides, P. incisa, P. αrpelata, P. prophylls grows into av egetative shoot or maximowiczii,P. mume and P. glandulosα. mixed shoot. That is,b uds in the axils of (b) Reproductive shoots abscising; vegeta- scalelike prophylls become new winter tive and mixed shoots persisting 四 buds. The species with this type is P. All reproductive shoots abscise and all g1iαyαnα. vegetative shoots persist. In addition,m ixed shoots also persist. That is,a ny shoots with (3) Shoot abscission leaves do not abscise. The species showing Species of Prunus usually abscise repro- this type is P. ogawana. ductive shoots by the end of the flower sea- (c) All shoots abscising son,w hile vegetative shoots persist. But All shoots,ir respective of the type,us ually extraordinary abscission has been reported in abscise by the winter of the shoot extension P. grayana. Our observations also revealed except the terminal vegetative shoots. As a variation in shoot abscission pattems in this result,t ree shape with as imple branching species. We recognized three types in this pattem is formed. The species exhibiting this character (Fig. 4). type is P. grayana. (a) Reproductive shoots abscising; vegeta- tive shoots persisting All reproductive shoots abscise by winter, 主 主 terminal shoot 、 Q 的 校 U 一 が・込 JF ・ぶ ~・..1 (a) (b) (c) Fig. 4. Shoot abscission in Prunus s. .1(a) Reproductive shoots abscise,wh ile vegetative shoots per- sist. (b) Mixed shoots and vegetative shoots persist. Reproductive shoots abscise. (c) All shoots except terminal ones abscise. Solid lined shoots persists and broken lined shoots abscise. Meaning of syrnbols esarne as those in Fig. 1. ぽ December 2001 Joumal of Japanese Botany Vol. 76 No. 6 335 Table 2. Result of observation on Prunus s. 1. in this study. Item 1: Types of winter bud growth. Item 2: Growth of buds in the axils of scale-like prophylls. Item 3: Shoot abscission. See the text for explana- tion of the Groups I-IV Subgenus Section Species Item 1 Item 2 Item 3 Group Cerasus Phy llomahaleb Prunus mωimowiczii a a a Pseudocerasus P. apetala a a a I P. cerasoides a a a I P. incisa a a a I P. jamasakura a a a - P. pendura a a a I a P. sargentii a a I a P. speciosa a a a I P. verecunda L a a - P.ogawana U b b H Microcerasus P. glandulosa C C a III Prunus P.mume a III C C Padus P. grayana d d C IV Discussion abscise,wh ile the shoots with leaves (mixed Grouping of observed species on the shoot and vegetative shoots) persist. orgamzatwn Group III [Prunus mume,P. glandulosa] We can categorize the observed species All winter buds develop into vegetative into four groups based on our observation of shoots and buds in the axils of scale-like the three types of shoot organization (Table prophylls develop into reproductive shoots 2). Detailed descriptions of each group are as simultaneously. All reproductive shoots ab- follows: scise and all vegetative shoots persist. Group 1[ Prunus jamαsakura,P. verecunda, Group IV [Prunus grayana] P. sargentii,P . speciosa,P . pendula,P . Mixed shoots and vegetative shoots de- cerαsoides, P. incisa, P. αrpetala, P. velop from winter buds. The winter buds are , maximowiczii] derived from smaller winter buds in the axils In these species the buds in the axils of of scale-like prophylls. All shoots except the leaves produce vegetative and reproductive terminal ones usually abscise before winter. shoots. Reproductive shoots abscise and vegetative shoots persist after flowering sea- Specific shoot orgamzatwn of Pmus son. Buds in the axils of scale-like prophylls ogawana are not present or,if present,th ey do not de- Prunus ogawana is basically similar to velop. other species of sect. Pseudocerasus and it Group II [Prunus ogawana] was difficult to find any distinction between All buds in the axils of leaves produce re- them except for the flowering of P. ogawana productive and mixed shoots (all shoots have at aj uvenile stage. For this reason,o n the flowers). The buds in the axil of scale-like basis of similar mo中hologyin the juv~nile prophylls in P. ogawana often develop into stage,M akino (1912),O hwi (1952) and vegetative shoots. All reproductive shoots Sugimoto (1961) treated it as af orm of P. 336 植物研究雑誌第76巻第6号 平成13年12月 jamasakura. Our study,h owever,r evealed of P. persica indicated that it has the same that P. ogawana is far different from the pattem (Guo et a .11992,K ervella et a1. other species in sect. Pseudocerasus in shoot 1995). Taxonomic studies based on morpho- organization (Table 2). logy have classified these three species into Wec onsider P. ogawana to show specific three different subgenera of Prunus s. ,.1but shoot organization because al1 winter buds the phylogenic analysis using DNA se- grow into shoots with flowers. Winter buds quences shows their close relationship. Our of other species of sect. Pseudocerasus study corroborates the results of that usually develop into shoots with no flowers phylogenic analysis,bu t the number of spe 閏 and shoots with some flowers and no leaves cies we analyzed is too few. Further analyses (reproductive) and some with just leaves of shoot organization in additional species in (vegetative); the vegetative shoot is photo- the subgenera Prunus, Amygdalus and synthetic and contributes to vegetative Cerasus sec .tMicrocerasus eneeded. 紅 growth. In P. ogawana,al l winter buds de- velop into shoots with flowers. If all shoots Shoot 01苫anizationof Prunus grayana had no leaves,as in the reproductive shoots Prunus grayana shows distinct shoot or- of other species of sect. Pseudocerasus,P . ganization. The most remarkable feature is ogawana would not develop leayes and the abscission of all shoots except the termi- could not. photosynthesize. It is understand- nal ones. In other species,al l shoots with able,t herefore,t hat P. ogawana develops leaves persist and the buds in the axils of leaves on reproductive shoots (= mixed their leaves develop into winter buds. In P. shoots) and that buds in the axils of scale- grayana,w e observed that the buds in the like prophylls develop into leafy shoots for axils of the leaves also expand and develop vegetative growth. some leaf primordia,b ut all shoots with those axillary buds abscise by the end of the Shoot organization of Group 1 -genus growing season. In other words,th e develop- Cerasus Group ment of buds in the leafaxils of non-terminal Prunus is sometimes divided into several shoots appe sto be of no use to the plant. 紅 genera (Komarov 1971,Y u et a .11986,O hba It is also ad istinct character of P. grayana 1992). Ohba (1992) united and raised subge- to develop mixed shoots as in P. ogawana, nus Cerasus sections Pseudocerasus and but,t he features of development are not the Phyllomahaleb to the genus Cerasus for the same. While P. ogawana develops mixed Japanese species. A recent phylogenetic shoots because of flowering in all shoots,P. analysis using DNA sequences also showed grayana develops vegetative shoots as well. the monophyly of Cerasus in the sense of We therefore consider that there may be Ohba (1992) (Lee and Wen 2001). In our some developmental differences in the for- study,al l species of subgenus Cerasus sec- mation process of mixed shoots between P. tions Pseudocerasus,ex cept P. ogawana,an d grayana and P. ogawana. Phyllomahaleb ein Group 1. ,紅 Further study Shoot organization of Group III -Prunus The present study indicates the specificity mume and P. glandulosa of shoot organization of P. ogawana. It is es- Our study showed that P. mume and P. pecially important that P. ogawanαdevelops glandulosa can be categorized into one cate- leaves on its mixed shoots since the develop- gory with the same shoot organization pat- ment of both flowers and leaves on the same tem. Previous studies on shoot organization shoot has never been observed in other spe- December 2001 Journal of Japanese Botany Vol. 76 No. 6 337 cies of genus Cerasus in the sense of Ohba mation in Prunus yedoensis Matsum. J. Jap. Soc. (1992). The leaf development process in this Hort. Sci. 58: 472-473. Guo Z.,G oi M.,T anaka M. and Chujo T. 1992. species is expected to be different from other Mo中hologicalstudies on the bud formation in species in subgenus Cerasus,w hich will be Prunus persica Batsch. Mem. Fac. Agric. Kagawa examined and discussed in af orthcoming Univ.44: 167-173. study. Kervella J.,P ages L. and Genard M. 1995. Growth Our results also indicate that shoot organi- context and fate ofaxillary meristems of young peach trees. lnfluence of pentshoot growth char- zation might be an important taxonomic fea- 訂 acteristic of emergence date. Ann. Bot. (Oxford). ture for reevaluating Prunus s. ,.1 since 76: 559-567. phylogenetic studies using DNA sequences Komarov V. L. 1971 Rosaceae: Rosideae,A mygdaloi- have basically agreed with our groupings of deae. Flora URSS 10: 1-512. English translation. species based on shoot organization. Smithonian lnstitution,W ashington,D C. Extensive observation of shoot organization Kubota H. 1993. Prunus ogawana and Prunus on additional species is needed. verecunda forma norioi. Kyoto Engei. 88: 7ー11. Lauri P. 1991. Donnees sur l'evoltion de la ramifica- Shoot organization in P. grayana also tion et de la floraison du pecher Prunus persica (L) needs further analysis. From an evolutionary Batsch. au cours de sa croissance. Ann. Sci. Na ,.t point of view,s hoot abscission following Bo .t11: 95-103. bud growth in this species is very inefficient. Lee S. and Wen 1. 2001. A phylogenetic analysis of It is difficult to understand how or why this Prunus and the Amygdaloideae (Rosaceae) using ITS sequences of nuclear ribosomal DNA. Amer. 1. pattern developed. Further ecological and Bo .t88: 150-160. evolutionary studies may provide an answer. Makino T. 1892. Notes on Japenese plants. Bot. Mag. Tokyo 16: 45-56 [mainly 52]. We thank Tama Forest Science Garden 一一一 1906.Observations on the flora of Japan. Bot. and Mr. T. Katsuki for providing of materi- Mag. Tokyo 20: 37-45[ mainly 44-45]. als. Wea lso thank Mr. T. Kurita and Mr. T. 一一一 1908.Observations on the flora of Japan. Bot. Mag. Tokyo 22: 93-102 [mainly 98-99]. 1wa yama for providing of materials and the 一一一 1912.Observations on the flora of Japan. Bot. advice how to propagate P. ogawana. We Mag. Tokyo 26: 172-184 [mainly 176-177]. acknowledge the general advice of Professor 一一一 1928.A contribution to the knowledge of the H. Ohba and Ms. A. Shimizu,th e University Flora of Japan. J. Jpn. Bot. 5: 11-14 [mainly 11- 12]. Museum of the University of Tokyo. Our Ohba H. 1989. Prunus. In: Satake Y.,H ara H.,W atari thanks also go to Dr. David E. Boufford, S. and Tominari T.,Wi ld Flowers of Japan,W oody Harverd University Herbaria for checking Plants 1: 186-198. Heibonsha Ltd.,T okyo. the English text and content. 一一一 1992.Jap anese cherry trees under the genus This study was partly supported by The Cerasus (Rosaceae). J. Jpn. Bot. 67: 276-281, New Technology Development Foundation, Ohwi J. 1952. A juvenile form of Prunus verecunda Koehne. J. Jpn. Bot. 27: 148. the Makino Botanical Garden, Kochi Rehder A. 1940. A manual of cultivated trees and Prefecture,a nd the public office of Sakawa- shurbs hardy in North America exclusive of the cho in Kochi Prefecture. subtropical and warmer temprate regions,2 nd ed. Macmillan,N ew Yo rk. References Sugimoto 1. 1961. New Keys of Japanese Trees. Rokugatsusha Ltd.,O saka. Ghora C. and Panigrahi G. 1995. The family Rosaceae Yu T.T.,Lu L.-T.,K u T.-C., Li c.-L. and Chen S.-X. in lndia,2 . Bishen Sigh Mahendra Pal Singh, ー 1986. Rosaceae (3), Amygdaloideae. Flora Dehra Dun. Reipublicae Popularis Sinicae,3 8. Science Press, Goi M.,G uo Z. and Tanaka M. 1989. Studies on the Beijing. bud formation in temperate ornamental tree and s耐ubs.2. Seasonal investigations on the bud for- 338 植物研究雑誌第76巻第6号 平成13年12月 岩元明敏,宮崎慶子 ,邑田 仁 .パラ科サクラ a b c 属のシュート構成の多様性およびワカキノサクラ の特異性について 番目のグループにはウメ及びニワザクラが, 4番 本研究ではパラ科サクラ属 (Prunus)植物のシュー 目のグループにはウワミズザクラがそれぞれ含ま ト構成の多様性を明らかにするため,日本に自生 れた.ワカキノサクラは他のサクラ節植物とは異 する種を中心にサクラ属13種(サクラ亜属サクラ なるグループに属し,全てのシュートに花をつけ 節10種, ミヤマザクラ節 1種,ユスラウメ節 1種, る,混成芽を持つ,前出葉肢芽が展開するなどシュー スモモ亜属 1種,ウワミズサクラ亜属 1種)につ ト構成に大きな特異性を持つことが分かつた.ま いて年間を通じた観察を行った.この結果,対象 た,グループ分けが基本的にサクラ属についての とした13種は (1)冬芽成長 (2)冬芽の鱗片化し 分子系統解析と一致することから,シュート構成 た前出葉肢芽の発達・伸長 (3)枝の脱落性とい がサクラ属内の系統関係を反映している可能性も う3つのシュート構成の特徴に注目することで, 示唆された. 明確に4つのグループに分けられ,多様性が確認 (a東京大学総合研究博物館, された.最初のグループには ワカキノサクラを 東京都立大学牧野標本館 b 除くサクラ亜属サクラ節とミヤマザクラ節の9種 現所属:慶応義塾高校講師, が, 2番目のグループにはワカキノサクラが, 3 東京大学大学院理学系研究科附属植物園) c

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