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Estrategia Reproductiva De Una Hierba Perenne: Hypoxis Decumbens (Hypoxidaceae)

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Estrategia Reproductiva De Una Hierba Perenne: Hypoxis Decumbens (Hypoxidaceae) Rev.Biol. Trop" 46(3):555-565, 1998 Estrategia reproductiva de una hierba perenne: Hypoxis decumbens (Hypoxidaceae) Elen�R¡¡jmllndez u. y Nelson RamírezI 1 Ce trode Botánica Tropical,Instituto de Biología Experimental,Universidad Central Ven!,!zuela,Aptdo. 471 14, � de . Caracas, . Venezuela. Fax 582 7535897. E-mai!: [email protected] Recibido 20-Il-1997.Corregido 19-V-1998. Aceptado I3-V-1998. AbStrae!: The r!!proouctive strategy of Hypoxisdecumbens L. (Hypoxidaceae), a perennial herb with hermaphroditic flowers,was studied tocheck the relations betweenreproductive system,life form, habitat, pollination system andother morphological and'ecological features. The. study wasdone in a secondarydeciduous forest with a population of 150- 200plants. Controlledpollinationswere done in twentyisolatedplant$.The �sults'pointtXI outthatH. decumbens ís an autogamousspecies, with outbreeding depression l!.pparently reflectirigenvironmental.adaptatíofl of the autogamous genotypes. The mainpollinators wete generalistinsects;Iike dipterans.IthasfunCtionall)' solitary andactinomorpbic flowers,with weH expose? floral parts, displayed ajJproximately at fue same heigl1t,and with inner andbefore �thesis dehiscence of the anthers, Thebiomassallocati()fi w�mainly to femalestruct�res, likefruits andseeds, wbich was . reflected in a low male-femalebiomass ratio, and in highfruit and seed seto The pollen-ovuleand the attr;lction-support biomass ralios were .Iow, associated with autogamous species, which have reliable pollination. The dispersal unit is rhe seeq ahdthe di$persal sffndromeis illyrmecochory, with tbeinfructesc!'!nce prostratt; 9vephe soil, the fmi! with grad­ ualdehiscence andwith lip\ddroP 1!'!tsandp lacental remains(also with Iipid.content) in the seedcoat.This species has several.structural .and functi0Il<llattributes that sustain a. mixéd mating system' and,nlthough the results indicate that most oftheprogenyis autoganmus; it does not sacrifice fueopportunity forout-crossing. K�y words: Hypoxys decumbell$,perennial herb,mixed mating system, autogamy,outbreedingdepressioni pollination, fmit andseed set,biomass allocation. Elpredominio deL.hermafroditismoentre las ción autógama que se presenta en las plantas co­ angiospermas (ca. 90% dy las poblaciones de sexuales se ve contrarrestada por la existencia plantas), sugiere la eXistenciade amplias venta­ de diferentes mecanÍsmos ql1e favorecen el en­ jas en reproducirse en forma paterna y materna trecruzamiento o xenogamia,como son los. sis­ simultáneamente (Lloyq1982, Richards 1990), temas de incompatibilidad, la dicogamia y la Entre éstas, la fadlidadde encontrarpareja en hercogatnia (Jain 1976, Lloyd .1982,Wyatt ambientes recientemente colonizados o cuando 1983, Richards 1990), Así, el 80% de las espe­ la dispersión del polen es ineficiente, y compar­ cíes hetinafroditáspresentasistemas de autoin­ tir el gasto en estructuras de atracción en plan­ compatibilidad, lo que lllsobliga a entrecruzar­ tas polinizadas por anímales (Sólblig1976).Sin separa obtenerdescendencía (Solbrig & Ro- embargo,laaparentefacilidadparalareproduc- 1Iins 1977). 556 REVISTA DE BIOL OGIA TROPICAL Estas dos posibilidades que se le presentan a decumbens, y determinarlas relaciones que se las plantas cosexuales, autofertilizarse o entre­ establecen entre el sistema reproductivo y la cruzarse, están asociadas con una gran variedad forma de vida, el ambiente, y los caracteres re­ de factores ecológicos, morfológicos y fisioló­ productivos. gicos que, combinados, determinan y caracteri­ zan el sistema reproductivo. Por ej emplo, el sis­ tema reproductivo guarda correlación con la es­ MATERIALESY MÉTODOS tabilidad ambiental. En los ambientes someti­ dos a continuas perturbaciones predominan las Área de estudio: El trabajo se desarrolló en especies autógamas, mientras que hay mayor el Arboretum de la Escuela de Biología de la proporción de xenogamía en los ambientes más Universidad Central de Venezuela (Colinas de estables (Opler et al. 1980). Por otra parte, las Bello Monte, Edo. Miranda; elevación aproxi­ plantas xenógamas muestran una mayor inver­ mada 1 100 ro s.n.m., 10° 30' N, 66° 53' W). El sión en estructuras masculinas puesto que de­ área presenta dos hectáreas de bosque deciduo ben tener un sistema eficiente de polinización, secundario en estado sucesional, donde existe que asegure la transferencia de polen, mientras una población de H. decumbens de unos 150- que las plantas autógamas pueden dirigir los re­ 200in dividuos que forman parte del estrato her­ cursos hacia estructuras femeninas (Charnov báceo junto con gramíneas y dicotiledóneas de 1979, Charlesworth & Charlesworth1981 , Lo­ muy bajo porte. vett Doust & Cavers 1982, Brunet 1992). Ade­ Biología floral: Se siguió la metodología más, la relación polen/óvulo, es menor a medi­ utilizada por Ramírez et al. (1990). Se determi­ da que es más eficiente la transferencia de polen nó la disposición y organización de las flores,el (Cruden 1977). Unido al costo de producción de tipo morfológico de la inflorescencia, la morfo­ polen está el costo de producción de estructuras logía floral, la estructura del perianto, el color de atracción de los polinizadores, que es menor floral principal y los colores adicionales de las a medida que aumenta el grado de autogamia partes, de acuerdo a Radford el al. (1974), Lep­ puesto que se reduce la dependencia de ellos. pik (1977) Y Küppers (1979). Se determinó la Todos los recursos no utilizados en atracción o orientaciónde las flores con respecto al sustra­ en función masculina pueden ser utilizados en to, y la longevidad floral, hora de antesis y pro­ producción de semillas, o función femenina ducción de néctar. La unidad de polinización se (Solbrig 1976, Schoen 1982, Cruden & Lyon caracterizó de acuerdo a la organización de la 1985, Brunet 1992, Antos & ABen 1994). Otra inflorescencia y al comportamiento de los poli­ asociación importante es entre forma de vida y nizadores. sistema reproductivo, estando la autogamia fre­ En diez floresen antesís, colectadas en plan­ cuentemente asociada a plantas de vida corta, tas diferentes, se determinó: diámetro de la flor como hierbas anuales, y la xenogamia a plantas (dos medidas perpendiculares), diámetro del perennes, tanto hierbas como arbustos y árboles áreareproductiva o áreacentral (donde se posan (Charlesworth 1992). los visitantes florales), altura de la flor (distan­ En Hypoxis decumbens L. (Hypoxidaceae) cia entre la base de la flory el ápice de la estruc­ se presenta una combinación de varios de estos tura floral más alta), distancia antera-estigma, factores ecológicos y morfológicos. Su forma tipo de dehiscencia de las anteras y tipo estig­ de vida es perenne, el hábitat en el que se desa­ mático (Radford et al. 1974). El áreaestígmáti­ rrolla está sometido a perturbaciones continuas, ca se estimó por observación en cámara lúcida y presenta flores hermafroditas, que pueden de 5 estigmas; paraesto se pesó un áreade pa­ permitir la autogamia pero también pueden pro­ pel conocida y se hizo la equivalencia con el pe­ mover la xenogamia. Basados en esta combina­ so del papel que representa el área estigmática, ción de factores, el objetivo del presente estudio tomando en cuenta el aumento utilizado en la fue caracterizarla estrategiareproductiva de H. cámara. '557 ·l Número y características del pOlén; En aurepeüriiZaci6ri¡ póliji�aitdo con ·polen de la estigmas colectados una vez cerradas la!¡'flótes propialflot. La PttrSeliCiáde �utogpnna se.deter- se determirló la.cmg¡Üle polen; los estigmas se minóenyertlas floráles no'maniptiladasy aisla- alcohút preservaron en 70% y luego se c010- das delltgtntes éxtemos hasta'la forriláción de rearon azul de se prepararon frutos. lla ptesenci§\9é agaihospermia se con algodón, pa- ' ' deter­ c . ra su observación al· mi roscopio ól'�iM;I\:Se · · minó en flores �ft(asculadas' ' en estado muy cóntó el número de' granos de polen sin '�)'iIii� . teinprano:de desiattÓllo'de la' yema. Posterior a (1cnary germinádos paraestimarelporcentaJé'de cada una de las�túeb�se verificóla fonriación . el . -c-·-�ertitiuación. ".'. ' . de frutosy serti\jllas vIábles� S'e'detern1lnó la forma, distribucióny orna- La proporción defrut6s'por flor y de seini- mentaciónde los g.·anos de polen. Se estim�.el Has poróvulo prodtioidaporp olinización cruza- c ¡íreaproyectada de los granos de polen coriáyu- da y aútopolinización� y poliniza ión automáti- ca da de unmicÍñmetré oculary el número degra� y autopolinización se cbmparó estadística- nos de polen·que puede cubrir el áreaes.tigmáti - mente por una prueba de G .(Sokal & Roblf �l , . ' 'ca. Elnúrr1�FO de granosde polen por flot'se es- 1981); si los valores de polinización cruzada y sigl;iíendO método Lloyd .'autopolinizacioo no di eren significativamente el déscJiÍtO'' pdr fi r�(::;b�\.,�;tixIl.ó " { :fu: . ; . ' �' . i, , , especie se considera autocompatible, y se �(l�?) : " , 'la rJiolbg4a,depo .... izaciáli: Sé':C�OnJos. considera autógamasi los válOre�deautopolini- >'1 fi1s�ct()g; que visitab.an 1�flQres;4pfm1te � pe- .'. zaéión y de polinizaci6n áutomáticitlio difieren " '4oc:ío�'de"(>bservaa'i()nde�7hara!í llno,cada en significativame4ité. Sé consider6 el valor de . 'U1l'promedl;o de .1$'fl�tes ��i\pén'tipd de óbSer� . 0,001 como límite/superior pataf':detérminar la . Lo� ifisect(j I O*'íiaron . derptobabilidad'aSociado ri'ya,dón . � cáPtullMós 'sé significm)ciad�:vU1,or ' " �f eS'fléCie.;8é�usaroti lQs;critérios r�� n�el9 al"eálGulo del�stáGístico G. t : c;,(i��isi�;¡tamaño del· apirlÍál�r�ál'gade!
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