mathématicien italien (Lugo 1853 - Bologne 1925). On lui doit la création du calcul différentiel absolu. Cet algorithme se trouve tout entier dans une remarque de Elwin Bruno Christoffel (1829-1900), mais ses méthodes ont leur raison d’être dans les rapports intimes qui les lient à la notion riemannienne de variété à n dimensions. Essentiellement attaché à la variété, cet algorithme est indépendant du choix des variables et se trouve être l’instrument naturel de toutes les recherches sur une telle variété, d’où son rôle fondamental en relativité générale.
J. I.
tentacule
Excroissance de la paroi du corps, plus ou moins longue, plus ou moins cylindrique, molle et mobile, utilisée soit simplement comme organe sensoriel, soit comme organe préhensile, éventuellement comme organe locomoteur.
Les tentacules n’existent que chez les Invertébrés (Arthropodes exceptés), bien que la trompe des Éléphants et les barbillons de certains Poissons répondent à la définition proposée.
Ces formations se rencontrent déjà chez certains Protozoaires ciliés, les Tentaculifères (ou Acinétiens), caractérisés précisément par la possession de fins prolongements creux terminés soit par une sorte de petite ventouse, soit par des pointes acérées. Ces tentacules sont utilisés pour la capture des proies et l’aspiration de leur contenu (suçoirs).
Parmi les Métazoaires, les tentacules les plus simples, uniquement sensoriels, se rencontrent chez les Mollusques gastropodes, où ils sont portés par la région antérieure du corps, généralement au nombre d’une paire. Chez les Gastropodes pulmonés stylommatophores (Escargot par exemple), il y en a deux paires, rétractiles, la postérieure portant des yeux à l’extrémité. Les Bivalves nageurs, comme la Coquille Saint-Jacques, sont dotés de nombreux petits tentacules simples sur le bord du manteau. Chez les Annélides polychètes errantes, les cirres parapodiaux, les antennes et les palpes prostomiens, les longs cirres péristomiens sont des tentacules sensoriels simples. Les Procordés (Ascidies, Amphioxus...) possèdent de petits tentacules prébuccaux qui, outre leur fonction sensorielle, jouent un rôle de filtration des particules alimentaires.
De très nombreux animaux aquatiques, surtout ceux qui sont fixés et de petite taille, se nourrissant de particules microscopiques (régime microphage), prennent leur nourriture grâce à un système complexe de tentacules ciliés qui entourent la bouche et jouent souvent en même temps le rôle d’organes respiratoires : le battement des cils détermine un courant d’eau qui est filtré par le rideau de tentacules ; les particules alimentaires sont agglutinées par du mucus et conduites vers la bouche par voie ciliaire (nutrition muco-ciliaire). L’ensemble des tentacules et de leur support est désigné sous le nom de lophophore. Le lophophore peut être profondément rétractile, comme chez les Bryozoaires, ou non, comme chez les Endoproctes ou les Brachiopodes ; chez ceux-ci, la base du lophophore est représentée par deux longs bras souvent enroulés en spirale, éventuellement soutenus par un squelette calcaire adhérent à la valve dorsale. Un dispositif de même type se trouve chez les Ptérobranches (Stomocordés), où le lophophore est constitué de 2 ou 12 bras garnis de longs tentacules ciliés, et chez les Pogonophores, dotés de très longs tentacules (de 1 à plus de 200) dont la face interne porte de nombreuses et fines pinnules. Chez ces animaux, dépourvus de tube digestif, les tentacules assurent non seulement la capture des particules alimentaires, mais leur digestion externe grâce à des enzymes sécrétées par des cellules situées à la base des pinnules ; celles-ci absorbent ensuite les produits de digestion comme le feraient des villosités intestinales. Notons que les Annélides polychètes tubicoles possèdent un panache de tentacules, souvent d’une grande beauté, qui fonctionne comme branchies en même temps qu’il filtre l’eau, retenant les particules alimentaires, qui sont conduites vers la bouche par voie muco-ciliaire.
Chez d’autres animaux microphages, les tentacules agglutinent les particules alimentaires grâce à du mucus et les portent à la bouche. C’est ainsi que fonctionnent les longs et fins « captacules » des Mollusques scaphopodes ou la couronne de tentacules péribuccaux des Holothuries (Concombres de mer). Chez ces Échinodermes, les tentacules (qui sont des ambulacres modifiés), simples et terminés par un petit disque ou ramifiés de façon plus ou moins compliquée, une fois chargés de particules alimentaires, sont invaginés dans le pharynx, qui récolte ces particules. Cela fait songer à l’enfant qui suce ses doigts maculés de confiture.
Mais les tentacules peuvent aussi intervenir dans la capture de proies plus ou moins volumineuses (nutrition macrophagique) ; ils présentent alors des adaptations particulières. C’est ainsi que, chez les Cnidaires, ils sont garnis de cellules hautement spécialisées — les cnidoblastes, dotés de minuscules harpons venimeux dévaginables très perfectionnés — et capables de capturer, immobiliser, voire tuer des proies parfois relativement très grosses, qu’ils amènent ensuite vers la bouche. Ces tentacules sont plus ou moins nombreux, parfois réduits à un seul, comme chez les polypes nourriciers (gastrozoïdes) des Siphonophores, où il est très long (filament pêcheur), doté de ramifications (tentilles) où les cnidoblastes sont très localisés (boutons urticants). Les tentacules des Hydres peuvent être utilisés dans un mode de locomotion par culbutes, l’animal se déplaçant alors en collant alternativement au substrat son disque pédieux et ses tentacules. Chez certaines Méduses, quelques tentacules très courts, ampullaires, sont transformés en organes sensoriels complexes (tactiles, olfactifs, statorécepteurs, photosensibles) : les rhopalies. Chez les Cténaires, il n’y a que deux tentacules, capables de se rétracter complètement dans deux profondes dépressions (gaines tentaculaires) de la paroi du corps. Longs et grêles, ils portent des tentilles riches en colloblastes, cellules montées sur une sorte de ressort à boudin et hérissées de globules adhésifs qui se collent aux proies. Les tentacules les plus spectaculaires du monde animal, utilisés comme organes sensoriels, de capture et de locomotion, sont certainement ceux des Mollusques céphalopodes, dibranchiaux plus particulièrement, où les 8 à 10 tentacules (ou bras), résultant de la modification de la région antérieure du pied, sont garnis, sur leur face interne, de nombreuses ventouses très perfectionnées.
J. Ch.
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