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Composite List of C4 Weeds

  • C. Dennis Elmore (a1) and Rex N. Paul (a1)


C4 plants account for a small fraction of the total number of plant species (fewer than 1000 out of 250 000). A larger proportion of the world's weed species possess C4 physiology. There are 2000 species in 500 genera and 125 families of flowering plants listed in the WSSA composite list of weeds. of that number, 146 species in 53 genera and 10 families exhibit the C4 syndrome. This, as a percentage, is 17-fold greater than the percentage of C4 plants in the total world plant population. In this report, we have listed the C4 -weed species and provide specific information concerning various aspects of their Kranz anatomy and C4 physiology.



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1. Baskin, J. M. and Baskin, C. C. 1981. Photosynthetic pathways indicated by leaf anatomy in fourteen summer annuals of cedar glades. Photosynthetica 15:205209.
2. Bender, M. M. and Smith, D. 1973. Classification of starch- and fructosan-accumulating grasses as C3 or C4 species by carbon isotope analysis. J. Br. Grassl. Soc. 28:97100.
3. Bjorkman, O. 1976. Adaptive and genetic aspects of C4 photosynthesis. Pages 287309 in Burris, R. H. and Black, C. C., eds. CO2 Metabolism and Plant Productivity. Univ. Park Press, Baltimore, MD.
4. Black, C. C. 1976. An assessment of C4 photosynthesis and productivity. Pages 397403 in Burris, R. H. and Black, C. C., eds. CO2 Metabolism and Plant Productivity. Univ. Park Press, Baltimore, MD.
5. Black, C. C., Chen, T. M., and Brown, R. H. 1969. Biochemical basis for plant competition. Weed Sci. 17:338344.
6. Brown, R. H. and Gracen, V. E. 1972. Distribution of the post-illumination CO2 burst among grasses. Crop Sci. 12:3033.
7. Brown, W. V. 1974. Another cytological difference among the Kranz subfamilies of the Gramineae. Bull. Torrey Bot. Club. 101:120124.
8. Brown, W. V. 1977. The Kranz syndrome and its subtypes in grass systematics. Mem. Torrey Bot. Club 23:197.
9. Carolin, R. C., Jacobs, S.W.L., and Vesk, M. 1977. The ultrastructure of Kranz cells in the family Cyperaceae. Bot. Gaz. 138:413419.
10. Chapman, K.S.R. and Hatch, M. D. 1981. Aspartate decarboxylation in bundle sheath cells of Zea mays and its possible contribution to C4 photosynthesis. Aust. J. Plant Physiol. 8:237248.
11. Crookston, R. K. and Moss, D. N. 1974. Interveinal distance for carbohydrate transport in leaves of C3 and C4 grasses. Crop Sci. 14:123125.
12. Downton, W.J.S. 1970. Preferential C4 -dicarboxylic acid synthesis, the post-illumination CO2 burst, carboxyl transfer step, and grana configurations in plants with C4 -photosynthesis. Can. J. Bot. 48:17951800.
13. Downton, W.J.S. 1975. The occurrence of C4 photosynthesis among plants. Photosynthetica 9:96105.
14. Edwards, G. E. and Huber, S. C. 1981. The C4 Pathway. Pages 237281 in Hatch, M. D. and Boardman, N. K., eds. The Biochemistry of Plants. Photosynthesis, Vol. 8. Academic Press, New York, NY.
15. Ellis, R. P. 1977. Distribution of the Kranz syndrome in the Southern African Eragrostoideae and Panicoideae according to bundle sheath anatomy and cytology. Agroplantae 9:73110.
16. Elmore, C. D. 1980. The paradox of no-correlation between leaf photosynthetic rates and crop yields. Pages 155167 in Hesketh, J. D. and Jones, J. W., eds. Predicting Photosynthesis for Ecosystem Models, Vol. II. CRC Press, Boca Raton, FL.
17. Furbank, R. T. and Badger, M. R. 1982. Photosynthetic oxygen exchange in attached leaves of C4 monocotyledons. Aust. J Plant Physiol. 9:553558.
18. Gifford, R. M. 1974. A comparison of potential photosynthesis, productivity and yield of plant species with differing photosynthetic metabolism. Aust. J. Plant Physiol. 1:107117.
19. Gutierrez, M., Edwards, G. E., and Brown, W. V. 1976. PEP carboxykinase containing species in the Brachiaria group of the subfamily Panicoideae. Biochem. Syst. Ecol. 4:4749.
20. Gutierrez, M., Gracen, W. E., and Edwards, G. E. 1974. Biochemical and cytological relationships in C4 plants. Planta 119:279300.
21. Gutierrez, M., Kanai, R., Huber, S. C., Ku, S. B., and Edwards, G. E. 1973. Photosynthesis in mesophyll protoplasts and bundle sheath cells of various types of C4 plants. I. Carboxylases and CO2 fixation studies. Z. Pflanzenphysiol. 72:305319.
22. Harrison, P. A. and Black, C. C. 1982. Two-dimensional electrophoretic mapping of proteins of bundle sheath and mesophyll cells of the C4 grass Digitaria sanguinalis (L.) Scop. (Grabgrass). Plant Physiol. 70:13591366.
23. Hatch, M. D. and Kagawa, T. 1974. Activity, location and role of NAD malic enzyme in leaves with C4 -pathway photosynthesis. Aust. J. Plant Physiol. 1:357369.
24. Hatch, M. D. and Kagawa, T. 1974. NAD malic enzyme in leaves with C4 pathway photosynthesis and its role in C4 acid decarboxylation. Arch. Biochem. Biophys. 160:346349.
25. Hattersley, P. W. 1982. 13C values of C4 types in grasses. Aust. J. Bot. 9:139154.
26. Hattersley, P. W. and Browning, A. J. 1981. Occurrence of the suberized lamella in leaves of grasses of different photosynthetic types. I. In parenchymatous bundle sheaths and PCR (“Kranz”) sheaths. Protoplasma 109:371401.
27. Hattersey, P. W. and Watson, L. 1976. C4 -grasses: An anatomical criterion for distinguishing between NADP-ME species and PCK or NAD-ME species. Aust. J. Bot. 14:297308.
28. Hegde, B. A. and Patil, T. M. 1981. Parthenium hysterophorus (L.), a C3 planst with “Kranz” syndrome. Photosynthetica 15:14.
29. Hesla, B. I., Tieszen, L. L., and Imbamba, S. K. 1982. A systematic survey of C3 and C4 photosynthesis in the Cyperaceae of Kenya, East Africa. Photosynthetica 16:196205.
30. Hnatiuk, R. J. 1980. C4 photosynthesis in the vegetation of Aldabra Atoll. Oecologia 44:327344.
31. Holm, L. G., Plucknett, D. L., Pancho, J. V., and Herberger, J. P. 1977. The World's Worst Weeds. Distribution and Biology. Univ. Press, Hawaii. 609.
32. Imbamba, S. K. and Papa, G. 1979. Distribution of the Kranz type anatomy in some dicotyledonous families of Kenya. Photosynthetica 13:315322.
33. Jenkins, C.L.D. 1983. A rational approach to the design of C4 specific herbicides. Abstr. Weed Sci. Soc. Am. 195.
34. Jimenez, M. S., Caballero-Ruano, A., and Gil, F. 1981. Atriplex glauca, a new plant with Kranz syndrome. Photosynthetica 15:2835.
35. Kennedy, R. A. and Laetsch, W. M. 1973. Relationship between leaf development and primary photosynthetic products in the C4 plant Portulaca oleracea L. Planta 115:113124.
36. Kennedy, R. A. and Laetsch, W. M. 1974. Plant species intermediate for C3, C4 photosynthesis. Science 184:10871089.
37. Koch, K. E. and Kennedy, R. A. 1982. Crassulacean acid metabolism in the succulent C4 dicot, Portulaca oleracea L. under natural environmental conditions. Plant Physiol. 69:757761.
38. Krenzer, E. G., Moss, D. N., and Crookston, R. K. 1975. Carbon dioxide compensation points of flowering plants. Plant Physiol. 56:194206.
39. Laetsch, W. M. 1974. The C4 syndrome: A structural analysis. Annu. Rev. Plant Physiol. 25:2752.
40. Long, S. P., Incoll, L. D., and Woolhouse, H. W. 1975. C4 photosynthesis in plants from cool temperate regions, with particular reference to Spartina townsendii . Nature (London) 257:622624.
41. Meinzer, F. C. 1978. Observations on the taxonomic and ecologic distribution of C4 photosynthesis in northwestern Central America. Rev. Biol. Trop. 26:359369.
42. Moss, D. N., Krenzer, E. G. Jr., and Brun, W. A. 1969. Carbon dioxide compensation points in related plant species. Science 164:187188.
43. Mulroy, T. W. and Rundel, P. W. 1977. Annual plants: Adaptations to desert environments. BioScience 27:109114.
44. Nakamoto, H., Ku, M.S.B., and Edwards, G. E. 1982. Inhibition of C4 photosynthesis by (benzomidooxy) acetic acid. Photosyn. Res. 3:293305.
45. Raghavendra, A. S. 1980. Characteristics of plant species intermediate between C3 and C4 pathways of photosynthesis: Their focus of mechanism and evolution of C4 syndrome. Photosynthetica 14:271283.
46. Raghavendra, A. S. and Das, V.S.R. 1976. Distribution of the C4 dicarboxylic acid pathway of photosynthesis in local monocotyledonous plants and its taxonomic significance. New Phytol. 76:301305.
47. Raghavendra, A. S. and Das, V.S.R. 1978. The occurrence of C4 -photosynthesis: A supplementary list of C4 plants reported during late 1974- and mid-1977. Photosynthetica 12:200208.
48. Rathnam, C.K.M., Raghavendra, A. S., and Das, V.S.R. 1976. Diversity in the arrangements of mesophyll cells among leaves of certain C4 dicotyledons in relation to C4 physiology. Z. Pflanzenphysiol. 77:283291.
49. Sankhla, N., Ziegler, H., Vyas, O. P., Stichler, W., and Trimborn, P. 1975. Eco-physiological studies on Indian Arid Zone plants. V. A screening of some species for the C4 -pathway of photosynthetic CO2 fixation. Oecologia 21:123129.
50. Sayre, R. T., Kennedy, R. A., and Pringnitz, D. J. 1979. Photosynthetic enzyme activities and localization in Mollugo verticillata populations differing in the levels of C3 and C4 cycle operation. Plant Physiol. 64:293299.
51. Smith, B. N. 1982. General characteristics of terrestrial plants (agronomic and forests) – C3, C4, and crassulacean acid metabolism plants. Pages 99118 in Mitsui, A. and Black, C. C. Jr., eds. CRC Handbook of Biosolar Resources. CRC Press, Boca Raton, FL.
52. Smith, B. N. and Brown, W. V. 1973. The Kranz syndrome in the Gramineae as indicated by carbon iostopic ratios. Am. J. Bot. 60:503513.
53. Smith, B. N. and Turner, B. L. 1975. Distribution of Kranz syndrome among Asteraceae. Am. J. Bot. 62:541545.
54. Szarek, S. R. 1979. The occurrence of crassulacean acid metabolism: A supplementary list during 1976 to 1979. Photosynthetica 13:467473.
55. Vogel, J. C. and Fuls, A. 1978. The geographical distribution of Kranz grasses in South Africa. S. Afr. J. Sci. 74:209215.
56. Waller, S. S. and Lewis, J. K. 1979. Occurrence of C3 and C4 photosynthetic pathways in North American Grasses. J. Range Mange. 32:1228.
57. Weed Science Society of America. 1971. Composite list of weeds. Weed Sci. 19:437476.
58. Welkie, G. W. and Caldwell, M. 1970. Leaf anatomy of species in some dicotyledon families as related to the C3 and C4 pathways of carbon fixation. Can. J. Bot. 48:21352146.
59. Ziegler, H., Batanouny, K. H., Sankhla, N., Vyas, O. P., and Stichler, W. 1981. The photosynthetic pathway types of some desert plants from India, Saudi Arabia, Egypt and Iraq. Oecologia 48:9399.


Composite List of C4 Weeds

  • C. Dennis Elmore (a1) and Rex N. Paul (a1)


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