Protoplasma 119, 219--221 (1984)
9 by Springer-Verlag1984
Occurrence of Crystals in Vascular Cambium
K. S. RAO and Y. S. DAVE*
Department of Biosciences, Sardar Patel University,
Vallabh Vidyanagar
Received June 9, 1983
Accepted July 6, 1983
Summary
Crystals are of common occurrencein plant tissues and considered to
be waste products resulting from active metabolism of plant cells.
They mostly occur in ray and verticalparenchyma of wood and bark
of angiosperm trees and often develop in tissues which soon cease to
be functional. While studying the annual rhythm of cambial activity
in two species of the Verbenaceae, Tectona grandis (RAo and DAVE
1981) and Gmelinaarborea (DAVEand RAO 1982), crystalsof calcium
oxalate have been observed in ray initials (Figs. 1-10). The crystals
were identified as calcium oxalate after a convincing chemical
behaviour towards dilute (10%) acetic, hydrochloric, and sulphuric
acids. They appear colorless when viewed with brightfield
illumination (Fig. 3), and show birefringence in polarized light
(Figs. 1 and 6). Radial, tangential, and transverse sections of 10-
15 ~tm thick were used to study the crystals in polarized light with a
Carl Zeiss Axiomat microscope.
In Tectona, crystals are grouped in more or less
spherical clusters about a central core which appears
non-crystalline. Such groups are known as druses
(Figs. 4 and 5), and seldom appear in the form o f a
fusiform shaped solitary unit (Fig.2, arrow). The
druses are often tetragonal and rarely polygonal in
sections. They vary in size and are often located singly
in the center or towards one end of cells. In Gmelina,
crystals are a sheaf of long acicular raphides and often
appear as complex conglomerates (Figs. 7 and 8).
Apparently, the density of crystals is high during the
dormant cambium period in both species. The size and
form of crystals differ from one cell to the other and
* Correspondence and Reprints: Department ofBiosciences, Sardar
Patel University, Vallabh Vidyanagar 388 120, India.
between the two species. However, the size is smaller
compared to those in vascular cells (Fig. 9). Crystals in
Gmelina are distributed on either side from ray initials
to phloem and xylem (Figs. 9 and 10) while in Tectona,
They are restricted only towards phloem (Figs. 2, 4,
and 5).
To date, there is no authentic information on
occurrence of crystals in cambial cells. During active
divisions meristematic cells mostly lack ergastic
inclusions (EsAu 1965). Appearance of crystals in
dormant and active ray initials of Tectona and Gmelina
cambium is enigmatic and not fully understood.
Since calcium oxalate crystals are observed directly
behind the cambium in parenchyma cells of angiosperm
wood, CHATTAWAY(1953) speculated that they are the
by-products of the active metabolism of the cambial
layer. KENNEDY et al. (1968) traced the progress of
crystal development from cambium in Abies grandis.
STEWART (1960) emphasized that calcium oxalate
crystals apparently are end result of detoxication
mechanism, whereby the potentially harmful oxalic
acid is rendered non-toxic by formation of insoluble
calcium salt. Based on these reports we consider that
the crystals in ray initials of Tectona and Gmelina are
the result of high metabolic rate generated by
physiologically active cambium with a consequent
accumulation in dormant cambium. However, a
comprehensive study of a greater number of plants is
needed to elucidate the genesis, growth, form,
distribution, and physiological role of crystals in
cambial cells. This m a y contribute significantly to the
taxonomy of these species.
220 K . S . RAo and Y. S. DAVE: Occurrence of Crystals in Vascular Cambium

Figs. 1-10
K. S. RAO and Y. S. DAVE: Occurrence of Crystals in Vascular Cambium 221

Abbreviations used in figures: C Z cambial zone, FI fusiform initials, P H phloem, R I ray initials, X Y xylem
Figs. 1-5. Crystals in ray cambial initials of Tectona grandis
Fig. 1. Tangential section showing crystal birefringence and distribution in cambial rays. x i05
Fig. 2. Transverse section showing fusiform type crystal (arrow) and druses in ray initials. Note crystal-free xylem ray cells, x 525
Fig. 3. Radial longitudinal section showing a druse (arrow) in a ray initial, x 820
Figs. 4 and 5. Radial longitudinal sections showing crystal distribution and form in ray initials. Xylem ray cells show birefringence with polarized
light. Fig. 4. x 290, Fig. 5. x 600
Figs. 6-10. Crystals in ray cambial initials of Gmelina arborea
Fig. 6. Tangential section showing birefringence and distribution of crystals in cambial rays. x 105
Fig. 7. Tangential section showing raphide bundles formed of many individuals of long needle-shaped crystals in a parallel arrangement in ray
initials, x 396
Fig. 8. Transverse section of cambial zone with the adjacent xylem and phloem showing the presence of raphide crystals evidently in ray initials.
The neighbouring fusiform cells are free of crystals, x 280
Fig. 9. Transverse section of xylem, cambium and phloem showing distribution of crystals in rays. x 85
Fig. 10. Radial longitudinal section showing crystals in ray initials. Note the small starch grains in xylem ray cells showing birefringence in
polarized light, x 310

All the figures were taken from unstained sections in polarized light except Figs. 3 and 8 which are from brightfield illumination and toluidine
blue stained section respectively to validate the presence of crystals in ray initials of cambium.

References KENNEDY, R. W., SASTRY,C. B. R., BARTON, G. M., 1968: Crystals in

CHATTAWAY,M. ~V[., 1953: The occurrence of heartwood crystals in
certain timbers. Aust. J. Bot. 1, 27--88.
DAVE, Y. S., RAO, K. S., 1982: Seasonal activity of vascular cambium
in Gmelina arborea Roxb, IAWA Bulletin 3, 59--65.
ESAU, K., 1965: Plant Anatomy, 2nd ed. New York: J. Wiley & Sons.
wood of genus Abies indigenous to Canada and the United States.
Can. J. Bot. 46, 1221--1228.
RAO, K. S., DAVE, Y. S., 1981: Seasonal variations in cambial
anatomy of Tectona grandis. Nord. J. Bot. 1, 535--542.
STEWART, C. M., 1960: Detoxication during secondary growth in
plants. Nature 186, 374--375.