Aerodynamics and Right-Left Symmetry in Wind Dispersal of Maple, Dipterocarps, Conifers and Some Genera of Apocyanaceae and Magnoliaceae
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J.Natn.Sci.Foundation Sri Lanka 2017 45 (3): 201-217 DOI: http://dx.doi.org/10.4038/jnsfsr.v45i3.8184 RESEARCH ARTICLE Aerodynamics and right-left symmetry in wind dispersal of maple, dipterocarps, conifers and some genera of apocyanaceae and magnoliaceae Kirthi Tennakone 6KHI¿HOG'ULYH3HDERG\0DVVDFKXVHWWV86$ Revised: 30 September 2016; Accepted: 19 January 2017 Abstract: 7KHZLQGGLVSHUVLQJVHHGVKDYHHYROYHGUHPDUNDEOH INTRODUCTION aerodynamic optimisation to minimise the speed of descent on GHWDFKPHQWIURPWKHWUHHVRWKDWWKH\FRXOGEHFDUULHGDZD\ E\ WKH ZLQG DQG GHSRVLWHG D GLVWDQFH DZD\ IURP WKH SDUHQW ,Q FHUWDLQ SODQWV ZLQG LV DQ HIIHFWLYH PRGH RI VHHG $QHI¿FLHQWSK\VLFDOPHFKDQLVPWKDWHQDEOHVVORZLQJGRZQ GLVSHUVDO 3ULRU WR HYROXWLRQ RI ELUGV ZLQG GLVSHUVDO could have been the sole mode of long distance seed WKHGHVFHQWLVURWDWLRQZKLFKUHGXFHVWKHWUDQVODWLRQDONLQHWLF HQHUJ\ 5RWDWLRQ DQG WUDQVODWLRQ DOZD\V GH¿QH D ULJKW RU GLVWULEXWLRQRQODQG&RQVHTXHQWO\WUHHVDGRSWLQJZLQG OHIWKDQGHGQHVV7KHSUHVHQWZRUNH[DPLQHVWKHDHURG\QDPLFV GLVSHUVDOKDYHHYROYHGVHHGVZLWKLQWULFDWHDHURG\QDPLF of the dispersal of seeds of many species in relation to DSSHQGDJHVWRVORZGRZQWKHGHVFHQWHQDEOLQJWKHPWREH ULJKWOHIWDV\PPHWU\JLYLQJH[DPSOHVWRVKRZWKDWLQQDWXUH WUDQVSRUWHGE\ZLQG *UHHQH 4XHVDGD3DQGRO¿ this symmetry is broken either spontaneously or intrinsically. ,]]R6WHYHQVRQ et al ., 2015). Plumes attached In the former situation, seeds have no geometrical right-left to lighter seeds achieve this via increase of the air drag. asymmetry, but an initial instability chooses one sense of +RZHYHUKHDYLHUVHHGVVWRULQJIRRGIRUWKHLQLWLDOSKDVH rotation (right or left). Whereas in the latter, geometrical of germination cannot utilise this method and the viable asymmetry dictates the sense of rotation. Seeds of the maple SK\VLFDODOWHUQDWLYHZRXOGEHWRPDNHWKHPVSLQZKLOH IDPLOL\EHORQJVWRWKH¿UVWFDWHJRU\DQGDWKHRUWLFDOPRGHOLV falling. Here the gravitational potential energy of a falling presented to explain the motion. Seeds of dipterocarps, conifers seed dissipates largely via transformation into rotational and some genera of apocyanaceae and magnoliaceae are found kinetic energy, effectively inducing an aerodynamic lift. WRSRVVHVVDKDQGHGQHVVZKLFKGHWHUPLQHVWKHURWDWLRQGXULQJ seed fall. In dipterocarps, conifers and magnoliaceae, both 7ZR RI WKH PRVW GLYHUVH DQG ZLGHO\ GLVWULEXWHG right-handed and left-handed seeds are produced in the same angiosperm tree genera are maple (Aceraceae) and tree but correlated to the handedness of the sprial phyllotaxy dipterocarps (Dipterocarpaceae). Former is one of the of the branch that bears the fruit. Apocyanaceae is found to PRVWFRPPRQWUHHVSHFLHVLQWHPSHUDWH]RQHVZKLOHWKH be unique and seeds of all genera have the same handedness. ODWWHU GRPLQDWHV UDLQ IRUHVWV LQ WKH WURSLFV $VKWRQ 7KH GHWDLOV RI H[WHQVLYH ¿HOG REVHUYDWLRQV H[SHULPHQWV *XQDWLOOHNH&RUOHWW 3ULPDFN*XQDWLOOHNH and theoretical interpretations are presented to illustrate the et al ., 2006; Lamarque, 2013). Their success is due to relationship of right-left asymmetry to the aerodynamics of bearing of auto-gyrating seeds. Pines and other conifers VHHGZLQGGLVSHUVDOSRLQWLQJRXWWKHLPSOLFDWLRQVRIHYROXWLQDU\ &RQLIHUDH GLVWULEXWHG ZRUOGZLGH ZHUH WKH GRPLQDQW optimisation in practical aerodynamics. WUHHVSHFLHVLQWKH0HVR]RLFHUD7KHQWKHLUSUHGRPLQDQW PRGHRIGLVSHUVDOPD\KDYHEHHQWKHZLQG 6WHYHQVRQ Keywords: Apocyanaceae, dipterocarps, magnoliaceae, maple, et al ., 2015), and present day conifer seeds carry a trait plant handedness, seed aerodynamics. for auto-gyration. [email protected] 202 Kirthi Tennakone Spinning and translational motion of an object involves WKH ¿UVW YDULHWLHV KDYH FORVH VLPLODULWLHV FRPSDUHG D KDQGHGQHVV 6RPHWKLQJ IDOOLQJ GRZQ DQG URWDWLQJ to the 5 th and 6 th WKDWDUHOLJKWHUDQGGLIIHULQZLQJDQG FORFNZLVH ZKHQ YLHZHG IURP DERYH GLIIHUV IURP WKH VHHG JHRPHWU\ 7KHUHIRUH PRVW H[SHULPHQWV ZHUH VDPH URWDWLQJ FRXQWHUFORFNZLVH $ PDSOH VDPDUD FRQGXFWHGZLWK1RUZD\PDSOHDQG%R[(OGHUVDPDUDV GURSSHGIURPDKHLJKWURWDWHVHLWKHUFORFNZLVHRUFRXQWHU $YHUDJH ZHLJKWV ZHUH GHWHUPLQHG IURP VDPSOHV FORFNZLVH $OWKRXJK WKHUH DUH UHSRUWV LQ OLWHUDWXUH DQG VDPSOHV ZHUH XVHG IRU GHWHUPLQDWLRQ RI WKH discussing kinematics of maple seed motion (Lentink GLPHQVLRQV:LQJVWUXFWXUHZDVH[DPLQHGE\ORZSRZHU et al ., 2009; Varshney et al ., 2012; Sohn et al. , 2014), to PLFURVFRS\ EHIRUH DQG DIWHU FKHPLFDO WUHDWPHQW ZLWK WKHNQRZOHGJHRIWKHDXWKRUWKHDVSHFWRIKDQGHGQHVV acetone and hypochlorite. These treatments enhanced the of motion has not been investigated. Observations WUDQVSDUHQF\RIZLQJVWRH[DPLQHWKHYHLQV7KHSRVLWLRQ indicate that maple samaras or any of the macroscopic RIWKHFHQWUHRIJUDYLW\ZDVDVVHVVHGE\EDODQFLQJWKH organs of maple trees have no handedness (right-left VDPDUD RQ D YHUWLFDOO\ SODFHG KRUL]RQWDO HGJH ERWK distinguishing property) attributing maple seed rotation OHQJWKZLVHDQGEUHDGWKZLVH3K\VLFDODOWHUDWLRQVWRWKH to a spontaneous parity violation (right-left symmetry). VDPDUDVZHUHPDGHE\FXWWLQJWKHZLQJVDQGVHHGHGJHV In contrast, the dipterocarp fruit and other organs of ZLWK VFLVVRUV VDQGLQJ WKH VHHG VXUIDFHV V\PPHWULFDOO\ their trees possess handedness and the sense of rotation RU DV\PPHWULFDOO\ DQG FRDWLQJ ZLWK SDUDI¿Q ZD[ DQG of the fruit during falling is predetermined. Dipterocarp SROLVKLQJ 7HUPLQDO YHORFLWLHV ZHUH GHWHUPLQHG E\ trees have distinct right-handed (RH) and left-handed releasing the seeds from varying heights and timing /+ WZLJVÀRZHUVDQGIUXLWV6WUDQJHO\WKHH[WHQVLYH WKHPRWLRQIRUWKH¿QDOPHWUHV7R¿[WKHVHQVH OLWHUDWXUHRQWD[RQRP\RIGLSWHURFDUSDFHDH )R[ZRUWK\ RIURWDWLRQ UHODWLRQWRWKHGRZQZDUGYHUWLFDO DUHJLRQ 1911; Ashton, 1980; Smitinand et al ., 1980; Simmathri LQWKHVHHGVXUIDFHZDVOLJKWO\SDLQWHG$VOHDGLQJHGJH 7XUQEXOO GRHV QRW UHFRUG REVHUYLQJ RI WKLV LV WKH ULGJH RYHU WKH ZLQJ REVHUYLQJ WKH XSSHUPRVW property. Pines and magnolias are also found to have VLGHHQDEOHV¿[LQJWKHVHQVHRIURWDWLRQ(DFKVHHGZDV KDQGHGQHVV DWWULEXWHV VLPLODU WR GLSWHURFDUSV ZKHUHDV dropped 100 times from a height of ~ 3 m to assess the the entire family of apocyanaceae is ambidextrous. probability of the either sense of rotation. The effect of WKH SRVLWLRQLQJ RI WKH VHHG RQ UHOHDVLQJ ZDV FDUHIXOO\ The investigation reported here based on extensive REVHUYHG %HIRUH DQG DIWHU HDFK SK\VLFDO PRGL¿FDWLRQ observations on symmetries in plants theoretically WKH DYHUDJH GLVWDQFH RI IDOO DW ZKLFK VSLQQLQJ DQDO\VHVWKHDHURG\QDPLFVRIZLQGGLVSHUVDOLQUHODWLRQ starts, (2) terminal velocity, and (3) sense of rotation to right-left symmetry. Detailed observation of the ZHUHDVVHVVHG7KHHIIHFWRIGU\LQJWKHVHHGDVDZKROH PRUSKRJHQHWLFIHDWXUHVRIVHHGVDVZHOODVSDUHQWWUHHV RUH[SRVLQJRQHVXUIDFHWRWKHVXQZDVDOVRH[DPLQHG DQGVLPSOHH[SHULPHQWDWLRQLOOXVWUDWHKRZHYROXWLRQKDV 7ZLQVLQRQHIUXLWZHUHH[DPLQHGWRGHWHUPLQHZKHWKHU RSWLPLVHGWKHZLQGGLVSHUVDOVWUDWHJ\ there exist any correlations of sense of rotation. METHODOLOGY Dipterocarpus zeylanicus VDPSOHV ZHUH FROOHFWHG IURP *DPSDKD .XUXQHJDOD .DOXWUD 0DWDUD DQG 7KH DQDO\VLV DQG GLVFXVVLRQ LV EDVHG RQ D ¿HOG .HJDOOH'LVWULFWVRI6UL/DQND7HUPLQDOYHORFLWLHVZHUH observations and sample collection, and experimentation DVFHUWDLQHGE\GURSSLQJWKHIUXLWVIURPPKHLJKW ZLWK ZLQG GLVSHUVLQJ VHHGV E H[DPLQDWLRQV RI The angular speeds of falling fruits through height of WKH RUJDQV RI SODQWV ÀRZHUV OHDI DQG EUDQFKLQJ aPZHUHHVWLPDWHGE\VWURERVFRSLFPHDVXUHPHQWV DUUDQJHPHQW WKDWEHDUWKHVHVHHGVWRDVFHUWDLQZKHWKHU :LQJ VXUIDFH ZDV H[DPLQHG E\ PDJQL¿FDWLRQ DQG WKH right-left asymmetries exist; and (c) experimentation DQJOHV EHWZHHQ WKH ORQJLWXGLQDO YHLQV ZHUH PHDVXUHG ZLWKSDSHUWR\PRGHOVVRWKDWSDUDPHWHUVFDQEHUHDGLO\ after imprinting the vein structure on paper by pressing adjusted. WKHLQNHGZLQJV7KHQXWZDVFXWLQWRWUDQVYHUVHVHFWLRQV to determine the geometry of sepal arrangement. Wing D :LQGGLVSHUVHGPDSOHVDPDUDVZHUHFROOHFWHGIURP DQGVHHGPRGL¿FDWLRQZDVGRQHLQWKHVDPHZD\DVIRU Peabody area, Massachusetts, United States. Seeds of maple. Sugar maple ( Acer saccharum 3D[ 1RUZD\ PDSOH (A. platanoides L.), Sycamore ( A. pseudoplatanus L.), (b) Whenever an opportunity arose, observations Red maple ( A. rubrum ), Silver maple ( A. saccharinum ZHUH FRQGXFWHG GXULQJ WKH SDVW \HDUV WR DVFHUWDLQ Pax) and Box Elder ( A. negundo L.) often mixed the handedness of the seeds and sense of spinning to XS ZHUH VHSDUDWHG DQG VWRUHG DW DPELHQW FRQGLWLRQV GHWHUPLQHZKHWKHUVHHGDV\PPHWULHVWUDQVFHQGWRRWKHU (~15 oC, relative humidity ~ 60 %). The samaras of RUJDQV 7KH DV\PPHWULHV LQ ÀRZHUV IUXLWV OHDYHV DQG September 2017 Journal of the National Science Foundation of Sri Lanka 45(3) Aerodynamics and right-left symmetry in seed wind dispersal 203 EUDQFKLQJ DUUDQJHPHQW ZHUH TXDOLWDWLYHO\ H[DPLQHG and dimensions of a maple samara varies from species 7KH UHODWLRQVKLS RI IUXLW KDQGHGQHVV WR WKDW RI ÀRZHU to species and perhaps climate conditions. Typically, a and leaf phyllotaxy in dipterocarps and related species 1RUZD\PDSOHVDPDUDKDVDOHQJWKaFPPD[LPXP ZDV H[WHQVLYHO\ LQYHVWLJDWHG$OPRVW DOO WKH JHQHUD RI EUHDGWKaFPZLWKDaPPVHHGDQGDPHGLDQZHLJKW DSRF\DQDFHDHLQ6UL/DQND ZLOGDQGRUQDPHQWDO ZHUH a PJ RI ZKLFK PRUH WKDQ a LV FRQWULEXWHG examined to identify traits of handedness in phyllotaxy, by the seed and the thick ridge of vein supporting the ÀRZHUVIUXLWVDQGVHHGV7ZR Sapu ( Michelia nilagrica ) ZLQJ )LJXUH 7KH%R[(OGHUVDPDUDVKDYHVPDOOHU YDULHWLHVJURZLQJLQWKH&HQWUDO3URYLQFHRI6UL/DQND dimensions; length ~ 3.8 cm, maximum breadth ~ 1 cm. DQG RUQDPHQWDO PDJQROLDV ZHUH XVHG WR DVFHUWDLQ 7KH\DUHDOVROLJKWHUZLWKDPHGLDQZHLJKWaPJZLWK WKH UHODWLRQVKLS RI OHDI SK\OORWD[\ WR ÀRZHU DQG IUXLW DQHORQJDWHGVHHGFDUU\LQJDERXWRIWKHZHLJKW7KH morphology.