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Twin Raphides in the Vitaceae and Araceae and a Model for Their Growth A Botany Publication and Papers Botany 9-1983 Twin Raphides in the Vitaceae and Araceae and a Model for Their Growth A. M. Cody Iowa State University Harry T. Horner Iowa State University, [email protected] Follow this and additional works at: http://lib.dr.iastate.edu/bot_pubs Part of the Botany Commons, Other Plant Sciences Commons, Plant Biology Commons, and the Plant Breeding and Genetics Commons Recommended Citation Cody, A. M. and Horner, Harry T., "Twin Raphides in the Vitaceae and Araceae and a Model for Their Growth" (1983). Botany Publication and Papers. 54. http://lib.dr.iastate.edu/bot_pubs/54 This Article is brought to you for free and open access by the Botany at Iowa State University Digital Repository. It has been accepted for inclusion in Botany Publication and Papers by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Twin Raphides in the Vitaceae and Araceae and a Model for Their Growth Abstract Several optical tests were employed to demonstrate twinning in raphides from 10 species in the family Vitaceae and one species in the Araceae. Scanning electron microscopy revealed that twin raphides differ from the reported shape of twin styloids and that raphides of the Vitaceae are "barbed" along two sides. A model is proposed to describe in vivo raphide production under saturated to supersaturated conditions of directional nutrient flow. Disciplines Botany | Other Plant Sciences | Plant Biology | Plant Breeding and Genetics Comments This article is from Botanical Gazette 144 (1983): 318. Posted with permission. This article is available at Iowa State University Digital Repository: http://lib.dr.iastate.edu/bot_pubs/54 Twin Raphides in the Vitaceae and Araceae and a Model for Their Growth Author(s): A. M. Cody and H. T. Horner Source: Botanical Gazette, Vol. 144, No. 3 (Sep., 1983), pp. 318-330 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/2474427 Accessed: 29-06-2016 19:23 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://about.jstor.org/terms JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to Botanical Gazette This content downloaded from 129.186.176.217 on Wed, 29 Jun 2016 19:23:54 UTC All use subject to http://about.jstor.org/terms BOT. GAz. 144(3):318-330. 1983. C 1983 by The University of Chicago. All rights reserved. 0006-807 1/83/4403-0016$02 .00 TWIN RAPHIDES IN THE VITACEAE AND ARACEAE AND A MODEL FOR THEIR GROWTH A. M. CODY AND H. T. HORNER Department of Botany, Iowa State University, Ames, Iowa 50011 Several optical tests were employed to demonstrate twinning in raphides from 10 species in the family Vitaceae and one species in the Araceae. Scanning electron microscopy revealed that twin raphides differ from the reported shape of twin styloids and that raphides of the Vitaceae are "barbed" along two sides. A model is proposed to describe in vivo raphide production under saturated to supersaturated conditions of directional nutrient flow. Introduction Planch., Rhoicissus capensis (Burm f.) Planch., Arrow-shaped raphides in the leaves and wood Tetrastigma voinierianum (Baltet) Pierre ex Gag- of various members of the grape family (Vitaceae) nep., Vitis vinifera var. Concordia, and Xantho- have been noted (SOLEREDER 1899; GARD 1900; soma sagittifolium (L.) Schott. were collected in the SOLEREDER and MEYER 1928; METCALFE and Bessey Hall Greenhouse; Parthenocissus quinque- CHALK 1950). ESAU (1965, figs. 9B, 33A) observed folia (L.) Planch., P. tricuspidata (Sieb. & Zucc.) the raphides of Vitis as four-sided in section. MET- Planch., P. vitacea (Knerr) Hitchc., and Vitis ri- CALFE and CHALK (1950) described leaf raphide paria Michx. were collected on the Iowa State Uni- sacs in the Vitaceae as "of variable lengths and versity Campus, Ames. Raphide crystals were often containing mucilage. the individual raph- extracted from fresh plant material so that the ef- ides [being] pointed at one end but bidentate at the fects of secondary physical or chemical alteration other in Cissus and Vitis." GARD (1900) found sim- of the crystals would be negligible. Raphides from ilar raphides in Ampelopsis and compared them the petioles were chosen for study because they with twin gypsum crystals collected at Montmarte, appeared to be largest and, therefore, more ame- France. nable to optical study. KOLLBECK, GOLDSCHMIDT, and SCHR6DER (1914) used optical analysis with polarized light to SCANNING ELECTRON MICROSCOPY (SEM) determine that arrow-shaped styloids from Iris Fresh longitudinal and transverse hand sections pseudacorus were twin crystals. FREY (1925) and of petioles were mounted on brass disks with con- ARNOTT (1981) suspected that forked raphides also ductive silver paste, coated with Au-Pd in a Po- might be twin crystals because they possessed a laron E5 100 sputter coating unit, and examined at dovetailed reentrant angle at one end of each crys- 15 kV with a JEOL JSM 35 SEM. tal and because of the apparent morphological sim- Individual raphides were extracted by dicing ilarity of raphides to Iris styloids. transversely cut petioles with a razor blade and Until we found unusually large raphides in the then touching the fragments to a piece of cover petioles of Cissus rotundifolia, the small size of glass. The crystals were allowed to settle for 1 min raphides and their complex optical behavior before the preparation was flooded with methanol, impeded an understanding of their crystallographic which was then drawn off with filter paper. This nature. Scanning electron microscopy, optical crys- procedure was repeated several times to remove tallographic techniques, and X-ray diffraction extraneous plant material. An examination of analysis were employed to study them and raphides methanol-washed and nonwashed crystals showed from nine other species in five genera of the family that the fine surface features of calcium oxalate Vitaceae and from Xanthosoma sagittifolium in the monohydrate (COM), which is insoluble in alcohol, family Araceae. were not affected by the washing. Cover glass frag- ments with crystals were mounted on brass disks Material and methods with silver conductive paste and sputter coated in SAMPLE COLLECTION the same manner as fresh petiole sections. To re- Cissus rotundifolia (Forsk.) Vahl., C. discolor duce SEM beam penetration, an operating voltage Blume, C. quadrangularis L., C. rhombifolia of 15 kV was used; in most instances, a sample tilt Manuscript received December 1982; revised manuscript re- of 500 was advantageous. ceived May 1983. OPTICAL MICROSCOPY Address for correspondence and reprints: Dr. H. T. HOR- NER, Department of Botany, Iowa State University, Ames, Single raphides were removed by macerating Iowa 50011. transversely cut petioles and touching them to mi- 318 This content downloaded from 129.186.176.217 on Wed, 29 Jun 2016 19:23:54 UTC All use subject to http://about.jstor.org/terms CODY & HORNER-TWIN RAPHIDES IN VITACEAE AND ARACEAE 319 croscope slides. After the crystals had settled, the peared perfectly planar along the entire crystal slide was rinsed with a pipetted stream of methanol length (figs. 6, 7), while the outer surfaces- of the to remove excess plant material. Raphide total crystal were irregular but in optical continuity with length and width at center point were measured at their twins. Because of the need for high magni- X 400 with a calibrated eyepiece. For each species, fication and the narrow width of the crystals, mea- 50 unbroken raphides were measured. A petro- surement of the angle between extinctions could graphic microscope with a rotating stage was used not be accurate. The values 15.30, 15.20, and 15.10 to examine raphides at X 500 in optical index oils were obtained for different crystals and were rea- in transmitted light and between crossed polariz- sonably close to the 14.70 accurately measured in ers. A quartz plate compensator was also used with large heart-shaped COM twins on (101) found in crossed polarizers to verify twinning. Bavarian coal mines (KOLLBECK et al. 1914). During rotation, the dark line of the twin plane X-RAY DIFFRACTION (fig. 8) moved across the width of the crystal, in- Entire petioles were processed in methanol in a dicating that the twin plane was being viewed at high-speed blender. Excess plant material was re- an angle rather than in normal cross section. The moved from the suspension with a separatory fun- lateral extent of movement was proportional to the nel where the heavier crystals settled to the bottom. deviation of the twin plane from vertical (fig. 9). Crystals were repeatedly suspended and allowed When a quartz plate compensator was used with to settle before most of the alcohol remaining with crossed polarizers, light passing through the twin the crystal fraction was evaporated. The mixtures was retarded to a greater extent when the principal were mounted on glass microscope slides and air- optical axis or direction of greatest light retardation dried. Removal of all plant material was not nec- of a crystal (double arrow, fig. 10) was aligned more essary. Samples were analyzed in a Phillips X-ray closely with the principal optical axis of the quartz diffractometer using CuKe radiation, 40 kV, 20 plate. Greater light retardation resulted in higher- mA, and 20/min scan speed between 100 and 520 order colors, so that one-half of the raphide ap- 20.
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