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Plant function and evolutionary biology
RESEARCH ARTICLE

Welwitschia mirabilis: CAM or not CAM — what is the answer?

Dieter J. von Willert A B , Nicole Armbrüster A , Tobias Drees A and Maik Zaborowski A
+ Author Affiliations
- Author Affiliations

A Institute for Plant Ecology, Westfälische Wilhelms-University, Hindenburgplatz 55, D-48143 Münster, Germany.

B Corresponding author. Email: Willert@uni-muenster.de

C This paper originates from a presentation at the IIIrd International Congress on Crassulacean Acid Metabolism, Cape Tribulation, Queensland, Australia, August 2001

Functional Plant Biology 32(5) 389-395 https://doi.org/10.1071/FP01241
Submitted: 19 November 2001  Accepted: 8 December 2004   Published: 27 May 2005

Abstract

After more than 20 years of extensive study we found clear evidence that Welwitschia mirabilis Hook.f. is able to take up CO2 at night in both of its natural ecosystems, the Namib desert and the Mopane savannah, and hence should be classified a crassulacean acid metabolism (CAM) plant. At six different sites, 85 W. mirabilis plants were marked and the growth rate of their leaves and leaf ribbons were measured over a period of 2.5 years. The slowest and the fastest growing plant of these 85 plants were from the Mopane savannah and from the north-west of the Brandberg massif, respectively. These were selected for the gas-exchange measurements of this study. Within the course of a year nocturnal CO2 uptake was found only in December and January when the nights were shortest and plants were flowering. CO2 uptake during the night was not pronounced and never accounted for more than 4% of the total CO2 uptake over 24 h. Maximum rates of nocturnal CO2 uptake never exceeded 0.1 µmol m–2 s–1 for the slowest and 0.2 µmol m–2 s–1 for the fastest growing plant. Neither water availability in the soil nor night temperature was found to determine nocturnal CO2 uptake in terms known for CAM plants. Regardless of the growing site all leaves of W. mirabilis contained high amounts of malic and citric acid. Small increases of acids over night as calculated from the gas exchange measurements are masked by the extremely uneven distribution of these acids in the leaves, making the feature of an overnight malic or citric acid accumulation an unsuited test for CAM in W. mirabilis. An increase in 13C discrimination with increasing distance from the coast was confirmed. Photorespiration was extremely high and followed air temperature around the leaf. Although the debate whether or not W. mirabilis is a CAM plant can be closed, no answer could be given why W. mirabilis makes so little use of CAM.

Keywords: citric acid, CO2 uptake, malic acid, Welwitschia mirabilis.


Acknowledgments

This investigation was partially sponsored by the Deutsche Forschungsgemeinschaft. We also thank Volkswagen (Uitenhage, RSA) for providing 4WD vehicles that allowed us to reach the plants with our heavy equipment even in rough field. AFROX (Windhoek) kindly supplied all gases needed for our measurements. We thank Frank von Willert for developing the necessary transfer, converting and calculating computer programmes. For unremitting help with the field work we thank F Bücker, U Wagner-Douglas, M Steinberg, A van der Merwe, J Olivier and M Veste. We are grateful to the Ministry of Environment and Tourism of Namibia for giving us the permission to conduct these investigations on Welwitschia mirabilis over a period of 20 years.


References


Bergmeyer, HU (1970). ‘Methoden der enzymatischen Analyse.’ (Verlag Chemie: Weinheim)

Cockburn W (1985) Variation in photosynthetic acid metabolism in vascular plants: CAM and related phenomena. New Phytologist 101, 3–24. open url image1

Dittrich, P ,  and  Huber, W (1974). Carbon dioxide metabolism in members of the Chlamydospermae. In ‘Proceedings of the 3rd international congress on photosynthesis’. pp. 1573–1578. (Elsevier: Amsterdam)

Eller BM, von Willert DJ, Brinckmann E, Baasch R (1983) Ecophysiological studies on Welwitschia mirabilis in the Namib desert. South African Journal of Botany 2, 209–223. open url image1

Gaff DF (1972) Drought resistance in Welwitschia mirabilis Hook. fil. Dinteria 7, 3–7. open url image1

Gillon JS, Griffiths H (1997) The influence of (photo)respiration on carbon isotope discrimination in plants. Plant, Cell and Environment 20, 1217–1230.
Crossref | GoogleScholarGoogle Scholar | open url image1

Griffiths, H , Borland, A , Gillon, J , Harwood, K , Maxwell, K ,  and  Wilson, J (1999). Stable isotopes reveal exchanges between soil, plants and atmosphere. In ‘Physiological plant ecology. The 39th symposium of the British Ecological Society. pp. 415–441. (Blackwell Science: Oxford)

Harris FS, Martin CE (1991) Correlation between CAM-cycling and photosynthetic gas exchange in five species of Talinum (Portulacaceae). Plant Physiology 96, 1118–1124. open url image1

Henschel JR, Seely MK (2000) Long-term patterns of Welwitschia mirabilis, a long lived plant of the Namib desert (including a bibliography). Plant Ecology 150, 7–26.
Crossref | GoogleScholarGoogle Scholar | open url image1

Herppich WB, Flach BM-T, von Willert DJ, Herppich M (1996) Field investigations of photosynthetic activity, gas exchange and water potential at different leaf ages of Welwitschia mirabilis during a severe drought. Flora 191, 59–66. open url image1

Herppich WB, Flach BM-T, von Willert DJ, Herppich M (1997) Field investigations in Welwitschia mirabilis during a severe drought. II. Influence of leaf age, leaf temperature and irradiance on photosynthesis and photoinhibition. Flora 192, 165–174. open url image1

Kozaki A, Takeba G (1996) Photorespiration protects C3 plants from photooxidation. Nature 384, 557–560.
Crossref | GoogleScholarGoogle Scholar | open url image1

Lancaster J, Lancaster N, Seely M (1984) Climate of the central Namib desert. Madoqua Series 14, 5–16. open url image1

Loeschen VS, Martin CE, Smith M, Eder SL (1993) Leaf anatomy and CO2 recycling during Crassulacean acid metabolism in twelve epiphytic species of Tillandsia (Bromeliaceae). International Journal of Plant Sciences 154, 100–106.
Crossref | GoogleScholarGoogle Scholar | open url image1

Martin CE, Highley M, Wang W-Z (1988) Ecophysiological significance of CO2 recycling via Crassulacean acid metabolism in Talinum calcycium Engelm. (Portulacaceae). Plant Physiology 86, 562–568. open url image1

Schulze ED, Schulze I (1976) Distribution and control of photosynthetic pathways in plants growing in the Namib desert, with special regard to Welwitschia mirabilis Hook. fil. Madoqua 9, 5–13. open url image1

Schulze E-D, Ziegler H, Stichler W (1976) Environmental control of Crassulacean acid metabolism in Welwitschia mirabilis Hook. fil in its range of natural distribution in the Namib desert. Oecologia 24, 323–334.
Crossref | GoogleScholarGoogle Scholar | open url image1

Smith BN, Epstein S (1971) Two categories of 13C / 12C ratios for higher plants. Plant Physiology 47, 380–384. open url image1

Szarek DR, Johnson B, Ting IP (1973) Drought adaptation in Opuntia basilaris: significance of recycling carbon through crassulacean acid metabolism. Plant Physiology 52, 539–541. open url image1

Ting IP (1985) Crassulacean acid metabolism. Annual Review of Plant Physiology 36, 595–662.
Crossref | GoogleScholarGoogle Scholar | open url image1

Ting IP, Burk JH (1983) Aspects of carbon metabolism in Welwitschia. Plant Science Letters 32, 279–285.
Crossref |
open url image1

Ting, IP ,  and  Rayder, L (1982). ‘Crassulacean acid metabolism.’ (American Society of Plant Physiologists: Rockville, MD)

von Caemmerer S, Farquhar GD (1981) Some relationships between the biochemistry of photosynthesis and the gas exchange of leaves. Planta 153, 376–387.
Crossref | GoogleScholarGoogle Scholar | open url image1

von Willert DJ (1985) Welwitschia mirabilis — new aspects in the biology of an old plant. Advances in Botanical Research 11, 157–191. open url image1

von Willert DJ (1994) Welwitschia mirabilis Hook. fil. — das Überlebenswunder der Namibwüste. Die Naturwissenschaften 81, 430–442.
Crossref | GoogleScholarGoogle Scholar | open url image1

von Willert DJ, Eller BM, Brinckmann E, Baasch R (1982) CO2 gas exchange and transpiration of Welwitschia mirabilis Hook.fil. in the central Namib desert. Oecologia 55, 21–29.
Crossref | GoogleScholarGoogle Scholar | open url image1

von Willert, DJ , Eller, BM , Werger, MJA , Brinckmann, E ,  and  Ihlenfeldt, H-D (1992). ‘Life strategies of succulents in deserts. With special reference to the Namib desert. Cambridge studies in ecology.’ (Cambridge University Press: Cambridge)

von Willert DJ, Wagner-Douglas U (1994) Water relations, CO2 exchange, water-use efficiency and growth of Welwitschia mirabilis Hook.fil. in three contrasting habitats of the Namib desert. Botanica Acta 107, 291–299. open url image1

Whatley JM (1975) The occurrence of a peripherical reticulum in plastids of the Gymnosperm, Welwitschia mirabilis.  New Phytologist 74, 215–220. open url image1

Winter K, Schramm MJ (1986) Analysis of stomatal and nonstomatal components in the environmental control of CO2 exchange in leaves of Welwitschia mirabilis.  Plant Physiology 82, 173–178. open url image1