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Calcium oxalate crystals in plants

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Abstract

Calcium (Ca) oxalate crystals occur in many plant species and in most organs and tissues. They generally form within cells although extracellular crystals have been reported. The crystal cells or idioblasts display ultrastructural modifications which are related to crystal precipitation. Crystal formation is usually associated with membranes, chambers, or inclusions found within the cell vacuole(s). Tubules, modified plastids and enlarged nuclei also have been reported in crystal idioblasts. The Ca oxalate crystals consist of either the monohydrate whewellite form, or the dihydrate weddellite form. A number of techniques exist for the identification of calcium oxalate. X-ray diffraction, Raman microprobe analysis and infrared spectroscopy are the most accurate. Many plant crystals assumed to be Ca oxalate have never been positively identified as such. In some instances, crystals have been classified as whewellite or weddellite solely on the basis of their shape. Certain evidence indicates that crystal shape may be independent of hydration form of Ca oxalate and that the vacuole crystal chamber membranes may act to mold crystal shape; however, the actual mechanism controlling shape is unknown.

Oxalic acid is formed via several major pathways. In plants, glycolate can be converted to oxalic acid. The oxidation occurs in two steps with glyoxylic acid as an intermediate and glycolic acid oxidase as the enzyme. Glyoxylic acid may be derived from enzymatic cleavage of isocitric acid. Oxaloacetate also can be split to form oxalate and acetate. Another significant precursor of oxalate in plants is L-ascorbic acid. The intermediate steps in the conversion of L-ascorbic acid to oxalate are not well defined. Oxalic acid formation in animals occurs by similar pathways and Ca oxalate crystals may be produced under certain conditions.

Various functions have been attributed to plant crystal idioblasts and crystals. There is evidence that oxalate synthesis is related to ionic balance. Plant crystals thus may be a manifestation of an effort to maintain an ionic equilibrium. In many plants oxalate is metabolized very slowly or not at all and is considered to be an end product of metabolism. Plant crystal idioblasts may function as a means of removing the oxalate which may otherwise accumulate in toxic quantities. Idioblast formation is dependent on the availability of both Ca and oxalate. Under Ca stress conditions, however, crystals may be reabsorbed indicating a storage function for the idioblasts for Ca. In addition, it has been suggested that the crystals serve purely as structural supports or as a protective device against foraging animals. The purpose of this review is to present an overview of plant crystal idioblasts and Ca oxalate crystals and to include the most recent literature.

Zusammenfassung

Calciumoxalat-Kristalle kommen in vielen Pflanzenarten und in fast allen Teilen und Geweben vor. Im allgemeinen werden sie innerhalb der Zellen gebildet, doch sind auch extrazelluläre Kristalle beschrieben. Die Kristallzellen oder Idioblasten zeigen besondere ultrastrukturelle Spezialitäten, die mit der Kristallbildung zusammenhängen. Diese ist meist mit Membranen, Räumen oder Einschlüssen in der order den Vakuole(n) verbunden. Tubuli, modifizierte Piastiden sowie vergrößerte Zellkerne sind ebenfalls für die Idioblasten erwähnt. Die Kristalle bestehen entweder aus der Monohydrat-(Whewellit) oder der Dihydrat-form (Weddellit). Mit verschiedenen Methoden können die Calciumoxalat-Kristalle identifiziert werden, die sichersten sind Röntgenstrahlen-Diffraction, Raman Mikroprobe-Analyse und die Infrarot-Spektroskopie. Viele pflanzliche Kristalle, die als Kalciumoxalat-Kristalle angesehen werden, sind nie eindeutig bestimmt worden. In einigen Fällen wurden sie lediglich auf Grund der Form als Whewellit oder Weddellit klassifiziert; es gibt aber Hinweise, dab die Kristallform unabhängig vom Hydratationsgrad ist und daß die Membranen der Kristallkammer in der Vakuole die Form beeinflussen. Der eigentliche Kontrollmechanismus ist noch unbekannt.

Oxalsäure wird auf verschiedenen Wegen in den Zellen synthetisiert, in Pflanzen kann Glycollat in Oxalsäure umgewandelt werden. Die Oxidation erfolgt in zwei Schritten mit Glyoxylsäure als Zwischenprodukt und dem Enzym Glycolsäure-Oxidase. Die Glyoxylasäure könnte durch enzymatische Spaltung der Isozitronensäure entstehen. Auch kann Oxalazetat in Oxalat und Azetat splitten. Eine weitere wichtige Ausgangssubstanz für Oxalat in Pflanzen ist die L-Ascorbin-säure, die Zwischenprodukte dieser Umbildung sind noch nicht eindeutig bekannt. Oxalsäure-Bildung erfolgt bei Tieren über ähnliche Stoffwechselwege und Calciumoxalat-Kristalle mögen in ähnlicher Weise entstehen.

Verschiedene Funktionen werden den pflanzlichen Kristall-Idioblasten und Kristallen zugeschrieben. Es besteht Gewißheit, daß die Oxalat-Synthese mit der Ionen-Balance korreliert ist; so können die pflanzlichen Kristalle die Anstrengungen belegen, ein Ionen-Gleichgewicht zu erhalten. In vielen Pflanzen wird das Oxalat nur in geringem Maße oder gar nicht weiter metabolisiert, es wird als Endprodukt des Stoffwechsels angesehen. Die Kristall-Idioblasten funktionieren wohl zur Entfernung des Oxalats, das sonst toxische Konzentration erreichen würde. Die Bildung hängt von der Verfügbarkeit sowohl des Calciums wie des Oxalats ab. Unter Calcium-Mangel kann es zur Reabsorption von Kristallen kommen, ein Hinweis auf eine gewisse Speicherfunktion der Idioblasten für Ca. Zusätzlich wird vermutet, daß die Kristalle mechanische Funktion haben und ein Schutz gegen fressende Tiere darstellen. Die Absicht dieses Review ist es, eine aktuelle Übersicht über die Kristall-Idioblasten und CaOxalat-Kristalle zu geben, die neueste Literatur berücksichtigt.

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  1. Vincent R. Franceschi

    Present address: Department of Botany, University of California, 95616, Davis, CA

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  1. Department of Botany, Iowa State University Ames, 50011, Iowa

    Vincent R. Franceschi & Harry T. Horner

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Franceschi, V.R., Horner, H.T. Calcium oxalate crystals in plants. Bot. Rev 46, 361–427 (1980). https://doi.org/10.1007/BF02860532

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