A Remarkable New Leaf Type With Unusual Photosynthetic Tissue in a Central Asiatic Genus of Chenopodiaceae

Abstract:

From the hygrohalophyte Borszczowia aralocaspica Bunge (Chenopodiaceae), a new leaf type with 1-layered chlorenchyma is described as „borszczovoid” and compared with other leaf types in subfamily Salsoloideae. The chlorenchyma is suspected to represent a unique C₄ type. Evidence is cited from anatomical studies and documented by micrographs and Carbon isotope determinations (ä¹³C values). The 1-layered photosynthetic tissue combines all essential anatomical characters of a 2-layered chlorenchyma of regular C₄ plants and is in intimate contact with concentrically arranged peripheral bundles. The ä¹³C values are − 13.03 ‰ from young stems and − 13.78 ‰ from leaves. The results are discussed in the anatomical, physiological and taxonomic framework. In addition, from distantly-related Suaeda species of section Conosperma the conospermoid leaf type is re-described. It is characterized by typical palisade and Kranz layers and differs from the C₄ suaedoid type by an external water-storaging hypodermis and an arrangement of Kranz cells reminiscent of the atriplicoid type from subfamily Chenopodioideae. From eight other species of Chenopodiaceae ä¹³C values are given for the first time.

Key words:

Borszczowia - Chenopodiaceae - Suaeda - C₄ plants - C₄ photosynthesis - Kranz anatomy - leaf types

References

• 01 Akhani,  H.,, Trimborn,  P.,, and Ziegler,  H.. (1997);  Photosynthetic pathways in Chenopodiaceae from Africa, Asia and Europe with their ecological, phytogeographical and taxonomical importance.  Pl. Syst. Evol.. 206 187-221
• 02 Bunge,  A.. (1878);  Salsolacearum novarum in Turkestania indigenarum descriptiones.  Trudy Imp. S.-Peterburgsk. Bot. Sada. 5 643-646
• 03 Butnik,  A.. (1984) Adaptaciya anatomicheskogo stroyeniya vidov semesistva Chenopodiaceae Vent. k aridnym usloviyam. Avtoreferat dissert. The adaptation of anatomical structure of the species of family Chenopodiaceae Vent. to arid conditions. Summary of Thesis. Tashkent; Akademiya Nauk Uzbekskoy SSR (FAN) 41 (in Russian)
• 04 Butnik,  A.. (1998) Possible pathways of evolution of mesophyll types in Chenopodiaceae Vent. Msc. Tashkent 12
• 05 Butnik,  A.,, Nigemanova,  R. N.,, Paiziyeva,  S. A.,, and Saidov,  D. K.. (1991) Ekologicheskaya anatomiya pustynnykh rastenii Srednei Azii l. Ecological anatomy of desert plants of Middle Asia. Tashkent; Akademiya Nauk Uzbekskoy SSR (FAN) 245 (in Russian)
• 06 Carolin,  R. C.,, Jacobs,  S. W. L.,, and Vesk,  M.. (1975);  Leaf structure in Chenopodiaceae.  Bot. Jahrb. Syst.. 95 226-255
• 07 Cerling,  Th. E.. (1999) Paleorecords of C₄ plants and ecosystems. C₄ Plant Biology. Sage, R. F. and Monson, R. K., eds. San Diego etc.; Academic Press 448-469
• 08 Dengler,  N. C., and Nelson,  T.. (1999) Leaf structure and development in C₄ plants. C₄ Plant Biology. Sage, R. F. and Monson, R. K., eds. San Diego etc.; Academic Press 133-172
• 09 Freitag,  H.,, Wucherer,  W.,, and Butnik,  A.. (1999) Leaf anatomy in Old World species of Suaeda (Chenopodiaceae): Taxonomic significance and evolutionary aspects. 14. Symposium Biodiversität und Evolutionsbiologie. Manitz, H. and Hellwig, F. H., eds. Jena; Institut für Spezielle Botanik 59
• 10 Gamaley,  Yu. V.. (1985);  Variacii kranc-anatomii u rastenenii pustyn' gobi i karakumy. Variations of Kranz-anatomy in plants of the Gobi and Kara-Kum.  Bot. Zh.. 70 1302-1313 (in Russian)
• 11 Gamaley,  Yu. V., and Shiredamba,  C.. (1988) Strukturnyye tipy pustynnykh rastenii (Biologicheskaya resursy i prirodnyye usloviya mongol'skoy narodnoy republiki 28). Structural types of desert plants. Leningrad; Nauka 45-66 (in Russian)
• 12 Hatch,  M. D.. (1999) C₄ photosynthesis: A historical overview. C₄ Plant Biology. Sage, R. F. and Monson, R. K., eds. San Diego etc.; Academic Press 17-46
• 13 Igersheim,  A., and Cichocki,  O.. (1996);  A simple method for microtome sectioning of prehistoric charcoal specimens, embedded in 2-hydroxyethyl methacrylate (HEMA).  Rev. Palaeobot. Palynol.. 92 389-393
• 14 Il'in,  M. M.. (1936) Chenopodiaceae. Flora S.S.S.R. 6. Moskva/Leningrad; Akad. Nauk 1-354
• 15 Kanai,  R., and Edwards,  G. E.. (1999) The biochemistry of C₄ photosynthesis. C₄ Plant Biology. Sage, R. F. and Monson, R. K., eds. San Diego etc.; Academic Press 449-487
• 16 Kulzer and Co. . (without year) Kulzer Histo-Technik. Ein neuartiges System zum Einbetten und Schneiden von histologischen Präparaten. Wehrheim; Kulzer and Co. 18
• 17 Leegood,  R. C., and Walker,  R. P.. (1999) Regulation of the C₄ pathway. C₄ Plant Biology. Sage, R. F. and Monson, R. K., eds. San Diego etc.; Academic Press 89-131
• 18 Monteil,  P.. (1906) Anatomic comparée de la feuille des chenopodiacées. Lons-le-Saunier; L. Declume 156
• 19 Osmond,  C. B.,, Winter,  K.,, and Ziegler,  H.. (1982);  Functional significance of different pathways of CO₂ fixation in photosynthesis.  Encycl. Plant Phys. N. S.. 12 B 479-547
• 20 Paulsen,  O.. (1911) Traek af vegetationen i Transkaspiens Lavland. Copenhagen; Gyldendal 239
• 21 P'yankov,  V. I.,, Kuz'min,  A. N.,, Demidov,  E. D.,, and Malsov,  A. I.. (1992);  Diversity of biochemical pathways of CO₂ fixation in plants of the families Poaceae and Chenopodiaceae from the arid zone of Central Asia.  Soviet Plant Phys.. 39 411-420
• 22 Sage,  R. W.,, Li,  M.,, and Monson,  R. K.. (1999) The taxonomic distribution of C₄ photosynthesis. C₄ Plant Biology. Sage, R. F. and Monson, R. K., eds. San Diego etc.; Academic Press 551-584
• 23 Volkens,  G.. (1887) Flora der ägyptisch-arabischen Wüste auf Grundlage anatomisch-physiologischer Forschungen. Berlin; Bornträger 156
• 24 Voznesenskaya,  E. V., and Gamaley,  Yu. V.. (1986);  Ul'trastrukturnaya kharakteristika list'yev s kranc-anatomiyey. The ultrastructural characteristics of leaf types with Kranz-anatomy.  Bot. Zh.. 71 1291-1307 (in Russian)