Abstract
Graniferous tracheary elements are unusual xylem conducting cells, characterized by having structural material in the lumen. They are known particularly from certain root parasitic angiosperms. The included material is usually granular but may also be amorphous or fibrillar, all having the same origin during differentiation of the tracheary element. Vessels and tracheids with such inclusions were first reported in 1895 by Heinricher inLathraea (Scrophulariaceae). During the early decades of this century graniferous tracheary elements were noted in a few other taxa by different workers but were largely forgotten until the early 1960’s. This paper reviews the early literature and the research carried out during the past twenty-five years on these peculiar cells. Graniferous tracheary elements are found typically in the body of the haustorium of the root parasite, especially in the expanded xylem tissue or “vascular core.” The cells are most widely documented for the hemi-parasitic Santalaceae and were first recorded there in 1910 by Benson. She named the cells “phloeotracheides,” believing they combined the functions of phloem and xylem conducting elements. Heinricher and Benson both considered the granules to be composed of amylodextrin starch and Benson also believed the cells contained an enucleated protoplast. Our work has demonstrated that the granules in the Santalaceae are proteinaceous and that the cells are dead at maturity. In 1978 we therefore renamed them “graniferous tracheary elements.” They occur in all species of the Santalaceae so far investigated and inAtkinsonia ligustrina andNuytsia floribunda of root parasitic Loranthaceae. In these two families graniferous tracheary elements have the same organization. Their occurrence in haustoria of root parasites from other families is also reviewed. Although few observations are yet available in the Olacaceae, the granules inXimenia americana are found to be starch grains, like those inLathraea, whereas those inOlax phyllanthi are protein. Such fundamentally different material in haustorial tracheary elements within the same taxonomic group naturally raises the question of relationships within the family. The function of graniferous tracheary elements has not been experimentally investigated but we have suggested that for some Santalaceae they might serve as a device for regulating the flow of xylem sap through the haustorium.
Zusammenfassung
Granulahaltige Xylem-Leitbahnen sind ungewöhnliche Xylem-Leitzellen, die durch strukturiertes Material im Lumen charakterisiert sind. Sie kommen besonders in gewissen wurzelparasitischen Angiospermen vor. Das eingeschlossene Material ist gewöhnlich körnig kann aber auch amorph oder fibrillär sein, da alle Typen den gleichen Ursprung während der Differenzierung der granulahaltigen Xylem-Leitbahnen haben. Gefässe und Tracheiden mit solchen Einschlüssen wurden zuerst von Heinricher (1895) inLathraea (Scrophulariaceae) erwähnt. Während der ersten Jahrzehnte dieses Jahrhunderts wurden granulahaltigen Xylem-Leitbahnen von verschiedenen Bearbeitern in einigen anderen Taxa bemerkt, gerieten aber weitgehend bis in die frühen 60er Jahre in Vergessenheit. Dieser Artikel gibt einen Überblick über die frühere Literatur und die Forschung über diese eigenartigen Zellen während der letzten 25 Jahre. Granulahaltigen Xylem-Leitbahnen sind nur im Haustorium-Körper von Wurzelparasiten, vor allem im erweiterten Xylemgewebe oder Tracheiden gefunden worden. Die Zellen sind am besten für die halbparasitischen Santalaceae dokumentiert und wurden an diesen zuerst von Benson (1910) erwähnt. Sie nannte die Zellen “phloeotracheides” in dem Glauben, dass sie die Funktion von Phloem und Xylem kombinierten. Heinricher und Benson betrachteten die Körner als aus Amylodextrin-Stärke bestehend und Benson glaubte auch, dass die Zellen einen kernlosen Protoplasten enthalten. Unsere Arbeit hat erwiesen, dass die Körner in den Santalaceae proteinhaltig und die ausgewachsenen Zellen abgestorben sind. In 1978 haben wir sie daher in granulahaltigen Xylem-Leitbahnen umbenannt. Sie kommen in allen Arten von bisher untersuchten Santalaceae und inAtkinsonia ligustrina undNuytsia floribunda aus der Familie Loranthaceae vor. In diesen zwei Familien besitzen die granulahaltigen Xylem-Leitbahnen die gleiche Organisation. Ihr Vorkommen in Haustorien von Wurzelparasiten aus anderen Familien wird ebenfalls besprochen. Obwohl bisher nur wenige Beschreibungen für die Olacaceae verfügbar sind, wurden die Körner inXimenia americana als Stärkekörner wie inLathraea erkannt, wogegen jene inOlax phyllanthi aus Protein bestehen. Solch grundlegend verschiedenes Material in den Tracheen-Elementen der Haustoria in der gleichen taxonomischen Gruppe wirft natürlich die Frage nach den Beziehungen innerhalb der Familie auf. Die Funktion der granulahaltigen Xylem-Leitbahnen ist nicht experimentell untersucht worden, aber wir haben für einige Santalaceae vorgeschlagen, dass sie als Mechanismus zur Regulierung des Xylemsaftflusses durch das Haustorium dienen.
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Fineran, B.A. Graniferous tracheary elements in haustoria of root parasitic angiosperms. Bot. Rev 51, 389–441 (1985). https://doi.org/10.1007/BF02860969
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DOI: https://doi.org/10.1007/BF02860969