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His¡¡work¡¡on¡¡climbing¡¡plants¡¡was¡¡his¡¡first¡¡sustained¡¡piece¡¡of¡¡work¡¡on¡¡the¡¡physiology¡¡of¡¡movement£»¡¡and¡¡he¡¡remarks¡¡in¡¡1864£º¡¡¡¡¡¨This¡¡has¡¡been¡¡new¡¡sort¡¡of¡¡work¡¡for¡¡me¡£¡¨¡¡¡¡£¨¡¨Life¡¡and¡¡Letters¡¨£»¡¡III¡£¡¡page¡¡315¡£¡¡¡¡He¡¡had£»¡¡however£»¡¡made¡¡a¡¡beginning¡¡on¡¡the¡¡movements¡¡of¡¡Drosera¡££©¡¡¡¡He¡¡goes¡¡on¡¡to¡¡remark¡¡with¡¡something¡¡of¡¡surprise£»¡¡¡¨I¡¡have¡¡been¡¡pleased¡¡to¡¡find¡¡what¡¡a¡¡capital¡¡guide¡¡for¡¡observations¡¡a¡¡full¡¡conviction¡¡of¡¡the¡¡change¡¡of¡¡species¡¡is¡£¡¨

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Thus¡¡he¡¡was¡¡led¡¡on¡¡to¡¡the¡¡belief¡¡that¡¡TWINING¡¡is¡¡the¡¡more¡¡ancient¡¡form¡¡of¡¡climbing£»¡¡and¡¡that¡¡tendril¡­climbers¡¡have¡¡been¡¡developed¡¡from¡¡twiners¡£¡¡¡¡In¡¡accordance¡¡with¡¡this¡¡view¡¡we¡¡find¡¡LEAF¡­CLIMBERS£»¡¡which¡¡may¡¡be¡¡looked¡¡on¡¡as¡¡incipient¡¡tendril¡­bearers£»¡¡occurring¡¡in¡¡the¡¡same¡¡genera¡¡with¡¡simple¡¡twiners¡£¡¡¡¡£¨Loc¡£¡¡cit¡£¡¡page¡¡195¡££©¡¡¡¡He¡¡called¡¡attention¡¡to¡¡the¡¡case¡¡of¡¡Maurandia¡¡semperflorens¡¡in¡¡which¡¡the¡¡young¡¡flower¡­stalks¡¡revolve¡¡spontaneously¡¡and¡¡are¡¡sensitive¡¡to¡¡a¡¡touch£»¡¡but¡¡neither¡¡of¡¡these¡¡qualities¡¡is¡¡of¡¡any¡¡perceptible¡¡value¡¡to¡¡the¡¡species¡£¡¡¡¡This¡¡forced¡¡him¡¡to¡¡believe¡¡that¡¡in¡¡other¡¡young¡¡plants¡¡the¡¡rudiments¡¡of¡¡the¡¡faculty¡¡needed¡¡for¡¡twining¡¡would¡¡be¡¡founda¡¡prophecy¡¡which¡¡he¡¡made¡¡good¡¡in¡¡his¡¡¡¨Power¡¡of¡¡Movement¡¨¡¡many¡¡years¡¡later¡£

In¡¡¡¨Climbing¡¡Plants¡¨¡¡he¡¡did¡¡little¡¡more¡¡than¡¡point¡¡out¡¡the¡¡remarkable¡¡fact¡¡that¡¡the¡¡habit¡¡of¡¡climbing¡¡is¡¡widely¡¡scattered¡¡through¡¡the¡¡vegetable¡¡kingdom¡£¡¡¡¡Thus¡¡climbers¡¡are¡¡to¡¡be¡¡found¡¡in¡¡35¡¡out¡¡of¡¡the¡¡59¡¡Phanerogamic¡¡Alliances¡¡of¡¡Lindley£»¡¡so¡¡that¡¡¡¨the¡¡conclusion¡¡is¡¡forced¡¡on¡¡our¡¡minds¡¡that¡¡the¡¡capacity¡¡of¡¡revolving¡¡£¨If¡¡a¡¡twining¡¡plant£»¡¡e¡£g¡£¡¡a¡¡hop£»¡¡is¡¡observed¡¡before¡¡it¡¡has¡¡begun¡¡to¡¡ascend¡¡a¡¡pole£»¡¡it¡¡will¡¡be¡¡noticed¡¡that£»¡¡owing¡¡to¡¡the¡¡curvature¡¡of¡¡the¡¡stem£»¡¡the¡¡tip¡¡is¡¡not¡¡vertical¡¡but¡¡hangs¡¡over¡¡in¡¡a¡¡roughly¡¡horizontal¡¡position¡£¡¡¡¡If¡¡such¡¡a¡¡shoot¡¡is¡¡watched¡¡it¡¡will¡¡be¡¡found¡¡that¡¡if£»¡¡for¡¡instance£»¡¡it¡¡points¡¡to¡¡the¡¡north¡¡at¡¡a¡¡given¡¡hour£»¡¡it¡¡will¡¡be¡¡found¡¡after¡¡a¡¡short¡¡interval¡¡pointing¡¡north¡­east£»¡¡then¡¡east£»¡¡and¡¡after¡¡about¡¡two¡¡hours¡¡it¡¡will¡¡once¡¡more¡¡be¡¡looking¡¡northward¡£¡¡¡¡The¡¡curvature¡¡of¡¡the¡¡stem¡¡depends¡¡on¡¡one¡¡side¡¡growing¡¡quicker¡¡than¡¡the¡¡opposite¡¡side£»¡¡and¡¡the¡¡revolving¡¡movement£»¡¡i¡£e¡£¡¡circumnutation£»¡¡depends¡¡on¡¡the¡¡region¡¡of¡¡quickest¡¡growth¡¡creeping¡¡gradually¡¡round¡¡the¡¡stem¡¡from¡¡south¡¡through¡¡west¡¡to¡¡south¡¡again¡£¡¡¡¡Other¡¡plants£»¡¡e¡£g¡£¡¡Phaseolus£»¡¡revolve¡¡in¡¡the¡¡opposite¡¡direction¡££©£»¡¡on¡¡which¡¡most¡¡climbers¡¡depend£»¡¡is¡¡inherent£»¡¡though¡¡undeveloped£»¡¡in¡¡almost¡¡every¡¡plant¡¡in¡¡the¡¡vegetable¡¡kingdom¡£¡¨¡¡¡¡£¨¡¨Climbing¡¡Plants¡¨£»¡¡page¡¡205¡££©

In¡¡the¡¡¡¨Origin¡¨¡¡£¨Edition¡¡I¡£¡¡page¡¡427£»¡¡Edition¡¡VI¡£¡¡page¡¡374¡££©¡¡Darwin¡¡speaks¡¡of¡¡the¡¡¡¨apparent¡¡paradox£»¡¡that¡¡the¡¡very¡¡same¡¡characters¡¡are¡¡analogical¡¡when¡¡one¡¡class¡¡or¡¡order¡¡is¡¡compared¡¡with¡¡another£»¡¡but¡¡give¡¡true¡¡affinities¡¡when¡¡the¡¡members¡¡of¡¡the¡¡same¡¡class¡¡or¡¡order¡¡are¡¡compared¡¡one¡¡with¡¡another¡£¡¨¡¡¡¡In¡¡this¡¡way¡¡we¡¡might¡¡perhaps¡¡say¡¡that¡¡the¡¡climbing¡¡of¡¡an¡¡ivy¡¡and¡¡a¡¡hop¡¡are¡¡analogical£»¡¡the¡¡resemblance¡¡depending¡¡on¡¡the¡¡adaptive¡¡result¡¡rather¡¡than¡¡on¡¡community¡¡of¡¡blood£»¡¡whereas¡¡the¡¡relation¡¡between¡¡a¡¡leaf¡­climber¡¡and¡¡a¡¡true¡¡tendril¡­bearer¡¡reveals¡¡descent¡£¡¡¡¡This¡¡particular¡¡resemblance¡¡was¡¡one¡¡in¡¡which¡¡my¡¡father¡¡took¡¡especial¡¡delight¡£¡¡¡¡He¡¡has¡¡described¡¡an¡¡interesting¡¡case¡¡occurring¡¡in¡¡the¡¡Fumariaceae¡£¡¡¡¡£¨¡¨Climbing¡¡Plants¡¨£»¡¡page¡¡195¡££©¡¡¡¡¡¨The¡¡terminal¡¡leaflets¡¡of¡¡the¡¡leaf¡­climbing¡¡Fumaria¡¡officinalis¡¡are¡¡not¡¡smaller¡¡than¡¡the¡¡other¡¡leaflets£»¡¡those¡¡of¡¡the¡¡leaf¡­climbing¡¡Adlumia¡¡cirrhosa¡¡are¡¡greatly¡¡reduced£»¡¡those¡¡of¡¡Corydalis¡¡claviculata¡¡£¨a¡¡plant¡¡which¡¡may¡¡be¡¡indifferently¡¡called¡¡a¡¡leaf¡­climber¡¡or¡¡a¡¡tendril¡­bearer£©¡¡are¡¡either¡¡reduced¡¡to¡¡microscopical¡¡dimensions¡¡or¡¡have¡¡their¡¡blades¡¡wholly¡¡aborted£»¡¡so¡¡that¡¡this¡¡plant¡¡is¡¡actually¡¡in¡¡a¡¡state¡¡of¡¡transition£»¡¡and¡¡finally¡¡in¡¡the¡¡Dicentra¡¡the¡¡tendrils¡¡are¡¡perfectly¡¡characterized¡£¡¨

It¡¡is¡¡a¡¡remarkable¡¡fact¡¡that¡¡the¡¡quality¡¡which£»¡¡broadly¡¡speaking£»¡¡forms¡¡the¡¡basis¡¡of¡¡the¡¡climbing¡¡habit¡¡£¨namely¡¡revolving¡¡nutation£»¡¡otherwise¡¡known¡¡as¡¡circumnutation£©¡¡subserves¡¡two¡¡distinct¡¡ends¡£¡¡¡¡One¡¡of¡¡these¡¡is¡¡the¡¡finding¡¡of¡¡a¡¡support£»¡¡and¡¡this¡¡is¡¡common¡¡to¡¡twiners¡¡and¡¡tendrils¡£¡¡¡¡Here¡¡the¡¡value¡¡ends¡¡as¡¡far¡¡as¡¡tendril¡­climbers¡¡are¡¡concerned£»¡¡but¡¡in¡¡twiners¡¡Darwin¡¡believed¡¡that¡¡the¡¡act¡¡of¡¡climbing¡¡round¡¡a¡¡support¡¡is¡¡a¡¡continuation¡¡of¡¡the¡¡revolving¡¡movement¡¡£¨circumnutation£©¡£¡¡¡¡If¡¡we¡¡imagine¡¡a¡¡man¡¡swinging¡¡a¡¡rope¡¡round¡¡his¡¡head¡¡and¡¡if¡¡we¡¡suppose¡¡the¡¡rope¡¡to¡¡strike¡¡a¡¡vertical¡¡post£»¡¡the¡¡free¡¡end¡¡will¡¡twine¡¡round¡¡it¡£¡¡¡¡This¡¡may¡¡serve¡¡as¡¡a¡¡rough¡¡model¡¡of¡¡twining¡¡as¡¡explained¡¡in¡¡the¡¡¡¨Movements¡¡and¡¡Habits¡¡of¡¡Climbing¡¡Plants¡¨¡£¡¡¡¡It¡¡is¡¡on¡¡these¡¡pointsthe¡¡nature¡¡of¡¡revolving¡¡nutation¡¡and¡¡the¡¡mechanism¡¡of¡¡twiningthat¡¡modern¡¡physiologists¡¡differ¡¡from¡¡Darwin¡£¡¡¡¡£¨See¡¡the¡¡discussion¡¡in¡¡Pfeffer's¡¡¡¨The¡¡Physiology¡¡of¡¡Plants¡¨¡¡Eng¡£¡¡Tr¡£¡¡£¨Oxford£»¡¡1906£©£»¡¡III¡£¡¡page¡¡34£»¡¡where¡¡the¡¡literature¡¡is¡¡given¡£¡¡¡¡Also¡¡Jost£»¡¡¡¨Vorlesungen¡¡uber¡¡Pflanzenphysiologie¡¨£»¡¡page¡¡562£»¡¡Jena£»¡¡1904¡££©

Their¡¡criticism¡¡originated¡¡in¡¡observations¡¡made¡¡on¡¡a¡¡revolving¡¡shoot¡¡which¡¡is¡¡removed¡¡from¡¡the¡¡action¡¡of¡¡gravity¡¡by¡¡keeping¡¡the¡¡plant¡¡slowly¡¡rotating¡¡about¡¡a¡¡horizontal¡¡axis¡¡by¡¡means¡¡of¡¡the¡¡instrument¡¡known¡¡as¡¡a¡¡klinostat¡£¡¡¡¡Under¡¡these¡¡conditions¡¡circumnutation¡¡becomes¡¡irregular¡¡or¡¡ceases¡¡altogether¡£¡¡¡¡When¡¡the¡¡same¡¡experiment¡¡is¡¡made¡¡with¡¡a¡¡plant¡¡which¡¡has¡¡twined¡¡spirally¡¡up¡¡a¡¡stick£»¡¡the¡¡process¡¡of¡¡climbing¡¡is¡¡checked¡¡and¡¡the¡¡last¡¡few¡¡turns¡¡become¡¡loosened¡¡or¡¡actually¡¡untwisted¡£¡¡¡¡From¡¡this¡¡it¡¡has¡¡been¡¡argued¡¡that¡¡Darwin¡¡was¡¡wrong¡¡in¡¡his¡¡description¡¡of¡¡circumnutation¡¡as¡¡an¡¡automatic¡¡change¡¡in¡¡the¡¡region¡¡of¡¡quickest¡¡growth¡£¡¡¡¡When¡¡the¡¡free¡¡end¡¡of¡¡a¡¡revolving¡¡shoot¡¡points¡¡towards¡¡the¡¡north¡¡there¡¡is¡¡no¡¡doubt¡¡that¡¡the¡¡south¡¡side¡¡has¡¡been¡¡elongating¡¡more¡¡than¡¡the¡¡north£»¡¡after¡¡a¡¡time¡¡it¡¡is¡¡plain¡¡from¡¡the¡¡shoot¡¡hanging¡¡over¡¡to¡¡the¡¡east¡¡that¡¡the¡¡west¡¡side¡¡of¡¡the¡¡plant¡¡has¡¡grown¡¡most£»¡¡and¡¡so¡¡on¡£¡¡¡¡This¡¡rhythmic¡¡change¡¡of¡¡the¡¡position¡¡of¡¡the¡¡region¡¡of¡¡greatest¡¡growth¡¡Darwin¡¡ascribes¡¡to¡¡an¡¡unknown¡¡internal¡¡regulating¡¡power¡£¡¡¡¡Some¡¡modern¡¡physiologists£»¡¡however£»¡¡attempt¡¡to¡¡explain¡¡the¡¡revolving¡¡movement¡¡as¡¡due¡¡to¡¡a¡¡particular¡¡form¡¡of¡¡sensitiveness¡¡to¡¡gravitation¡¡which¡¡it¡¡is¡¡not¡¡necessary¡¡to¡¡discuss¡¡in¡¡detail¡¡in¡¡this¡¡place¡£¡¡¡¡It¡¡is¡¡sufficient¡¡for¡¡my¡¡purpose¡¡to¡¡point¡¡out¡¡that¡¡Darwin's¡¡explanation¡¡of¡¡circumnutation¡¡is¡¡not¡¡universally¡¡accepted¡£¡¡¡¡Personally¡¡I¡¡believe¡¡that¡¡circumnutation¡¡is¡¡automaticis¡¡primarily¡¡due¡¡to¡¡internal¡¡stimuli¡£¡¡¡¡It¡¡is¡¡however¡¡in¡¡some¡¡way¡¡connected¡¡with¡¡gravitational¡¡sensitiveness£»¡¡since¡¡the¡¡movement¡¡normally¡¡occurs¡¡round¡¡a¡¡vertical¡¡line¡£¡¡¡¡It¡¡is¡¡not¡¡unnatural¡¡that£»¡¡when¡¡the¡¡plant¡¡has¡¡no¡¡external¡¡stimulus¡¡by¡¡which¡¡the¡¡vertical¡¡can¡¡be¡¡recognised£»¡¡the¡¡revolving¡¡movement¡¡should¡¡be¡¡upset¡£

Very¡¡much¡¡the¡¡same¡¡may¡¡be¡¡said¡¡of¡¡the¡¡act¡¡of¡¡twining£»¡¡namely¡¡that¡¡most¡¡physiologists¡¡refuse¡¡to¡¡accept¡¡Darwin's¡¡view¡¡£¨above¡¡referred¡¡to£©¡¡that¡¡twining¡¡is¡¡the¡¡direct¡¡result¡¡of¡¡circumnutation¡£¡¡¡¡Everyone¡¡must¡¡allow¡¡that¡¡the¡¡two¡¡phenomena¡¡are¡¡in¡¡some¡¡way¡¡connected£»¡¡since¡¡a¡¡plant¡¡which¡¡circumnutates¡¡clockwise£»¡¡i¡£e¡£¡¡with¡¡the¡¡sun£»¡¡twines¡¡in¡¡the¡¡same¡¡direction£»¡¡and¡¡vice¡¡versa¡£¡¡¡¡It¡¡must¡¡also¡¡be¡¡granted¡¡that¡¡geotropism¡¡has¡¡a¡¡bearing¡¡on¡¡the¡¡problem£»¡¡since¡¡all¡¡plants¡¡twine¡¡upwards£»¡¡and¡¡cannot¡¡twine¡¡along¡¡a¡¡horizontal¡¡support¡£¡¡¡¡But¡¡how¡¡these¡¡two¡¡factors¡¡are¡¡combined£»¡¡and¡¡whether¡¡any¡¡£¨
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