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semillas la vida en cápsulas de tiempo - Clh.es

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The force of the b<strong>la</strong>st catapults the f<strong>la</strong>t discoid seeds, of which there are as many as there are carpels, as far as 14m.The name sandbox tree dat<strong>es</strong> back to the time before the inv<strong>en</strong>tion of blotting paper and fountain p<strong>en</strong>s, wh<strong>en</strong> thefruits served as containers for the sand used to blot the ink that ran profusely from goose quills.Active explosiv<strong>es</strong>Fl<strong>es</strong>hy <strong>de</strong>hisc<strong>en</strong>t fruits are able to build up suffici<strong>en</strong>t pr<strong>es</strong>sure to explo<strong>de</strong> by means of tissu<strong>es</strong> of thin-walled cells,which increase their internal cell pr<strong>es</strong>sure (called turgor) by taking up additional water. As the tissu<strong>es</strong> swell, at somepoint the pr<strong>es</strong>sure against neighbouring ine<strong>la</strong>stic <strong>la</strong>yers is so high that the slight<strong>es</strong>t movem<strong>en</strong>t of the fruit sets offthe explosion that expels the seeds. A passing animal can trigger the mechanism and although the seeds of suchfruits are g<strong>en</strong>erally smooth and non-sticky, they may become <strong>en</strong>tangled in its fur and be carried some distance.C<strong>la</strong>ssic exampl<strong>es</strong> of actively exploding fruits are the touch-me-not (Impati<strong>en</strong>s spp.) in the balsam family(Balsaminaceae), and the squirting cucumber (Ecballium e<strong>la</strong>terium) in the gourd family (Cucurbitaceae).Anyone who has p<strong>la</strong>yed with the fruits of the touch-me-not, g<strong>en</strong>tly squeezing them betwe<strong>en</strong> two fingers,knows that its name is appropriate. Its p<strong>en</strong>dulous fruits are composed of five carpels, with pre-formed <strong>de</strong>hisc<strong>en</strong>celin<strong>es</strong> betwe<strong>en</strong> them. Pr<strong>es</strong>sure builds up in the outer <strong>la</strong>yers of the pericarp and wh<strong>en</strong> it is strong <strong>en</strong>ough to separatethe carpels, the slight<strong>es</strong>t movem<strong>en</strong>t caus<strong>es</strong> the fruits to explo<strong>de</strong>. The outsi<strong>de</strong> of the pericarp expands more thanthe insi<strong>de</strong> causing the fruit valv<strong>es</strong> to curve inwards in a lightning-fast movem<strong>en</strong>t that hurls the seeds up to 5maway. The trigger for the explosion can be a passing animal, raindrops or water dripping off a tree, the wind, orev<strong>en</strong> seeds <strong>la</strong>unched from a neighbouring fruit. As the fruits of the Mediterranean squirting cucumber rip<strong>en</strong>, theinner tissue <strong>la</strong>yers build up turgor pr<strong>es</strong>sure against the thick, rather ine<strong>la</strong>stic outer skin. The pre-<strong>de</strong>terminedbreaking line at the base of the stalk points up in the air as the fruit turns almost 180º against the pedicel.Ev<strong>en</strong>tually, the slight<strong>es</strong>t movem<strong>en</strong>t caus<strong>es</strong> the stalk to pop out like a cork, leaving a hole through which bothwatery juice and seeds squirt out. The pr<strong>es</strong>sure insi<strong>de</strong> the fruit is so high that the seeds can travel more than 10m.The true record breakers are dwarf mistleto<strong>es</strong>, which apply more or l<strong>es</strong>s the same principle as the squirtingcucumber. Predominantly native to North America, they are parasit<strong>es</strong> on pine tre<strong>es</strong> and belong to the g<strong>en</strong>usArceuthobium in the Christmas mistletoe family (Viscaceae; rec<strong>en</strong>tly united with the sandalwood family, Santa<strong>la</strong>ceae).Whereas other mistleto<strong>es</strong> rely upon birds for the dispersal of their seeds, dwarf mistleto<strong>es</strong> (with the exception ofArceuthobium verticilliflorum) have evolved explosively <strong>de</strong>hisc<strong>en</strong>t fruits. Just as in the squirting cucumber, the pedicelb<strong>en</strong>ds downwards as the fruit matur<strong>es</strong> and a <strong>de</strong>hisc<strong>en</strong>ce line forms around its point of attachm<strong>en</strong>t. Once the fruithas reached maturity, the slight<strong>es</strong>t touch dislodg<strong>es</strong> it from the pedicel. The pr<strong>es</strong>sure that builds up insi<strong>de</strong> the darkgre<strong>en</strong>, single-see<strong>de</strong>d berry is so high that it fir<strong>es</strong> the tiny sticky seed through the hole in the rubbery fruit wall overa distance of up to 16m at the remarkable speed of 2m per second (97km per hour). A single pon<strong>de</strong>rosa pineinfected with Arceuthobium campylopodum may be bombar<strong>de</strong>d by more than two million seeds, which overwinter onthe surface where they <strong>la</strong>nd and ger-minate the following spring. If a seed is lucky <strong>en</strong>ough to <strong>la</strong>nd on a compatiblehost tree, its embryo grows, supported by the photosynthetic <strong>en</strong>dosperm tissue, and <strong>en</strong>ters the host’s bark.Aceuthobium speci<strong>es</strong> can live insi<strong>de</strong> their host for months or ev<strong>en</strong> years before producing a p<strong>la</strong>nt.Whether the seeds are catapulted from passively or actively exploding fruits, they are dispersed over nomore than a few metr<strong>es</strong>. The advantage of ballistic dispersal is that it is cheap, requiring no animal reward andusually very little in terms of specialised structur<strong>es</strong>. The preferred strategy of annual p<strong>la</strong>nts is to disperse theirseeds in a way that allows them to maintain and expand an existing popu<strong>la</strong>tion in the same p<strong>la</strong>ce rather thancolonising new territori<strong>es</strong>. For per<strong>en</strong>nial p<strong>la</strong>nts the most important factor is that the seeds should avoidcompetition with the par<strong>en</strong>t p<strong>la</strong>nt. However, the ejection of the seeds from their fruits is oft<strong>en</strong> only the first phaseof their dispersal history. Many seeds, such as the squirting cucumber, pansi<strong>es</strong>, spurg<strong>es</strong> and gorse, have an <strong>en</strong>ergyrich,oily app<strong>en</strong>dage, which functions as an edible bait that lur<strong>es</strong> ants into carrying them away from the par<strong>en</strong>tp<strong>la</strong>nt. Apart from such app<strong>en</strong>dag<strong>es</strong>, ballistically dispersed seeds are usually small, smooth and more or l<strong>es</strong>sspherical, <strong>en</strong>suring low air r<strong>es</strong>istance.Geocarpy – or how do peanuts <strong>en</strong>d up un<strong>de</strong>rground?This is a legitimate qu<strong>es</strong>tion if one has never se<strong>en</strong> a peanut p<strong>la</strong>nt (Arachis hypogaea, Fabaceae). After all, peanuts arefruits and fruits <strong>de</strong>velop from flowers, and flowers need to be pollinated, which mostly happ<strong>en</strong>s in the air. So howcan a fruit <strong>en</strong>d up un<strong>de</strong>rground? The answer is that the p<strong>la</strong>nt itself buri<strong>es</strong> them. After pollination and fertilization, theovary is pushed down into the ground, h<strong>en</strong>ce the name geocarpy for this type of self-dispersal. The single carpel of thepeanut flower sits at the <strong>en</strong>d of a specialised stalk-like organ called the gynophore. As soon as the ovul<strong>es</strong> are fertilised,the gynophore b<strong>en</strong>ds down and elongat<strong>es</strong> until it has pushed the young, pointed ovary un<strong>de</strong>rground. Once the ovaryhas reached its subterranean <strong>de</strong>stination, it swells to produce peanuts. Other legum<strong>es</strong> that bury their fruits in the sameway are the Bambara groundnut from W<strong>es</strong>t Africa (Vigna subterranea) and Astragalus hypogaeus from w<strong>es</strong>t Siberia.Geocarpy is mostly found in annual p<strong>la</strong>nts living in dry, hot climat<strong>es</strong> such as <strong>de</strong>serts, grass<strong>la</strong>nds and savannahhabitats. Here, burying the fruits is not only a safeguard against grazing animals, it primarily <strong>en</strong>sur<strong>es</strong> that the nextg<strong>en</strong>eration is kept in a suitable location in an inhospitable <strong>en</strong>vironm<strong>en</strong>t.One rare temperate example of a p<strong>la</strong>nt that buri<strong>es</strong> its fruits is the ivy-leafed toadf<strong>la</strong>x (Cymba<strong>la</strong>ria muralis,P<strong>la</strong>ntaginaceae), a common wild flower on walls and rocks in Britain, Europe and North America. Wh<strong>en</strong> inbloom, the flowers face the sun. As soon as they have be<strong>en</strong> pollinated, their pedicels turn away from the sunlightand seek out sha<strong>de</strong> by growing longer and searching the substrate for suitably dark cracks and cranni<strong>es</strong> in whichto <strong>de</strong>posit their fruit. Wh<strong>en</strong> ripe, the capsul<strong>es</strong> op<strong>en</strong> to release a number of small, irregu<strong>la</strong>rly ornam<strong>en</strong>ted seeds.Careful p<strong>la</strong>nting by the mother p<strong>la</strong>nt and the rugged surface of the fruits prev<strong>en</strong>t them from rolling out of theirprotected <strong>en</strong>vironm<strong>en</strong>t.Self-burying drillsSome diaspor<strong>es</strong> bury themselv<strong>es</strong> after dispersal. Apart from keeping them hidd<strong>en</strong> from animal predators, this selfburialis regar<strong>de</strong>d as an adaptation to dry soils as it allows the diaspore to reach the more humid <strong>la</strong>yers just un<strong>de</strong>rthe soil surface. The rotating movem<strong>en</strong>t of their hygroscopic app<strong>en</strong>dag<strong>es</strong>, which twist and untwist with chang<strong>es</strong>in humidity, allows the diaspore to drill itself into the ground. This behaviour is most famously found in speci<strong>es</strong>of stork’s bills, e.g. red-stem stork’s bill (Erodium cicutarium), and musky stork’s bill (E. moschatum). Their diaspor<strong>es</strong>consist of fragm<strong>en</strong>ts (fruitlets) of a schizocarpic fruit with a long beak. Once the fruit has come apart, each singl<strong>es</strong>ee<strong>de</strong>dfruitlet retains a share of the beak, which serv<strong>es</strong> as a hygroscopically moving awn. The same mechanismis found in a number of speci<strong>es</strong> of the very distantly re<strong>la</strong>ted grass family (Poaceae). The florets of wild oat (Av<strong>en</strong>afatua), wild barley (Hor<strong>de</strong>um vulgare ssp. spontaneum) and the needle-and-thread grass (Stipa comata) are equippedwith an awn, the basal portion of which twists and untwists with chang<strong>es</strong> in moisture levels. The drilling floretsof the Australian corkscrew spear-grass (Stipa setacea) are so sharp that they are able to p<strong>en</strong>etrate the wool andskin of sheep. Once embed<strong>de</strong>d in the fl<strong>es</strong>h of the animal, the curved hairs on the surface of the florets make themalmost impossible to dislodge. The muscle movem<strong>en</strong>ts of the tortured animals drag the fruits <strong>de</strong>eper and <strong>de</strong>eperinto their body. Florets of Stipa setacea have allegedly be<strong>en</strong> found in the heart muscle of <strong>de</strong>ad sheep.Creeps and jerksHygroscopically moving app<strong>en</strong>dag<strong>es</strong> <strong>en</strong>able other diaspor<strong>es</strong> to creep along the ground for short distanc<strong>es</strong>. Suchcreeping diaspor<strong>es</strong> are found in some grass<strong>es</strong>, and in the sunflower family (Asteraceae) and teasel family(Dipsacaceae). Their fruits have a hygroscopically moving pappus (modified calyx). For example, the fruits(cypse<strong>la</strong>s) of the cornflower (C<strong>en</strong>taurea cyanus, Asteraceae) are crowned by a tuft of short, stiff, scale-like pappussegm<strong>en</strong>ts, which are too small to p<strong>la</strong>y any role in wind dispersal. Their function is rather differ<strong>en</strong>t: with changinghumidity the pappus scal<strong>es</strong> repeatedly move in and out, thus pushing the fruits a few c<strong>en</strong>timetr<strong>es</strong> over theground. The very short, forward-pointing teeth along their margins prev<strong>en</strong>t movem<strong>en</strong>t in the opposite direction.The distanc<strong>es</strong> they creep are short but at least they move away from the par<strong>en</strong>t p<strong>la</strong>nt; wind and rainwater maycarry them further. A more targeted additional strategy has giv<strong>en</strong> them an edible swelling at the base tospecifically attract ants, a mo<strong>de</strong> of dispersal that will be discussed in <strong>de</strong>tail.Specialised awns <strong>en</strong>able the fruits of certain grass<strong>es</strong> (e.g. Arrh<strong>en</strong>atherum e<strong>la</strong>tius, Av<strong>en</strong>a sterilis) to jerk and jumpfor short distanc<strong>es</strong>. The distal part of their long-kneed awns is straight whereas the lower part is helically twistedand extremely hygroscopic. With changing humidity the basal part winds or unwinds, turning the straight distalpart of the awn. Since each fruit has two awns, which turn in opposite directions, their distal parts ev<strong>en</strong>tuallymeet and become <strong>en</strong>tangled. The t<strong>en</strong>sion that builds up betwe<strong>en</strong> them is released wh<strong>en</strong> the pr<strong>es</strong>sure is strong<strong>en</strong>ough to push the distal parts past each other. Within a split second, the jerky movem<strong>en</strong>t of the awns catapultsthe diaspore into the air.Dispersal by animalsAnimal movem<strong>en</strong>ts are l<strong>es</strong>s haphazard than wind and water making animals much more reliable as dispersa<strong>la</strong>g<strong>en</strong>ts. A p<strong>la</strong>nt that manag<strong>es</strong> to <strong>de</strong>velop a re<strong>la</strong>tionship with animals needs fewer seeds to guarantee the survivalof the speci<strong>es</strong>. The evolution of animal-dispersed diaspor<strong>es</strong> clearly has many advantag<strong>es</strong> and so it is not surprisingthat some fifty per c<strong>en</strong>t of gymnosperms (Ephedra, Gnetum, Ginkgo, a few conifers and all cycads) use animals toassist in the dispersal of their seeds. Once again it was the angiosperms that perfected the shift from abiotic tobiotic dispersal ag<strong>en</strong>ts by evolving a fascinating spectrum of strategi<strong>es</strong> <strong>en</strong>abling them to travel either on or insi<strong>de</strong>animals. Th<strong>es</strong>e close, sometim<strong>es</strong> highly specialised re<strong>la</strong>tionships betwe<strong>en</strong> angiosperms and animal dispersersprovi<strong>de</strong> another key to the un<strong>de</strong>rstanding of the evolutionary succ<strong>es</strong>s of this group.The evid<strong>en</strong>ce of such strategi<strong>es</strong> in p<strong>la</strong>nts li<strong>es</strong> in every sweet, juicy fruit we <strong>en</strong>joy. The sweet pulp of thefruit is a bait to lure pot<strong>en</strong>tial dispersers into swallowing the seeds and dispersing them in their faec<strong>es</strong>. If they arefortunate, the seeds will <strong>la</strong>nd in a suitable p<strong>la</strong>ce to grow well away from the shadow of the mother p<strong>la</strong>nt. Thisform of dispersal is called <strong>en</strong>dozoochory, “dispersal insi<strong>de</strong> an animal”. Endozoochory is a frequ<strong>en</strong>tly <strong>en</strong>counteredph<strong>en</strong>om<strong>en</strong>on in many famili<strong>es</strong> with fl<strong>es</strong>hy fruits, e.g. Ericaceae, Rosaceae, So<strong>la</strong>naceae. Endozoochorouslydispersed seeds do not pr<strong>es</strong><strong>en</strong>t any conspicuous adaptations to facilitate their dispersal. They are usually smooth,globose to ovoid, and either covered in a hard <strong>en</strong>docarp (if borne in drup<strong>es</strong>) or with a hard seed coat (if borneEnglish texts 281

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