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The Requirement for Sodium As a Micronutrient by Species Having the C4 Dicarboxylic Photosynthetic Pathway

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The Requirement for Sodium As a Micronutrient by Species Having the C4 Dicarboxylic Photosynthetic Pathway Plant Physiol. (1972) 49, 794-797 The Requirement for Sodium as a Micronutrient by Species Having the C4 Dicarboxylic Photosynthetic Pathway Received for publication September 28, 1971 P. F. BROWNELL AND C. J. CROSSLAND Department of Botany, James Cook University of North Queensland, Townsville, Queensland, Australia ABSTRACT MATERIALS AND METHODS Six species having characteristics of plants with the C4 di- Seeds of Echinochloa litilis L. Ohwi et Yabuno (Japanese carboxylic photosynthetic pathway, Echinochloa utilis L. Ohwi millet), Cynodon dactylon L. (Bermuda grass), Poa pratensis et Yabuno (Japanese millet), Cynodon dactylon L. (Bermuda L. (Kentucky blue grass). Ainaranthus tricolor L. cv. Early grass), Kyllinga brevifolia Rottb., Amaranthus tricolor L. cv. splendour, Kochia childsii Hort., and Portulaca grandiflora Early splendour, Kochia childsii Hort., and Portulaca grandi- Hook (rose moss) were obtained from commercial seed sup- flora Hook (rose moss), responded decisively to 0.1 mili- pliers. Seeds of Kyllinga brevifolia Rottb. were collected lo- equivalent per liter NaCl supplied to their culture solutions cally. The procedures for germination and growth of plants initially containing less than 0.08 microequivalent per liter Na. under conditions of low sodium have been described previously Chlorosis and necrosis occurred in leaves of plants not receiv- (2). The basal culture solution had the following composition ing sodium. Portulaca failed to set flower in the sodium-de- expressed in ftmoles/liter: KNO3, 5,000: Ca(NOj). 4,000; ficient cultures. Under similar conditions Poa pratensis L. (Kentucky blue grass) having characteristics of the C3 pho- Table I. Responises of Variouis Plaiits to Sodium tosynthetic pathway made normal growth and did not respond to the addition of sodium. It is concluded from these results and Yield Age Lesions in Signifi- previously reported work that sodium is generally essential for at Plants Not cance but not for with the Species' lIar- Receiving No 0.1 meq of Dif- species having the C4 pathway species C3 vest Sodium addi- liter ference pathway. tion ANaCl days g dry wt/plait % Gramineae Poa pratensis L. (Keni- 52 None 0.0236 0. 0206 NS2 tucky blue grass) Echilnochloa utilis L. Ohwi 22 Chlorosis and 0.404 1 0.713 0.1 et Yabuno (Japanese necrosis millet) Cy,iodooi dactylon L. 47 Chlorosis 0.178 0. 337 Sodium was first shown to be essential for the blue-green (Bermuda grass) alga Anabaena cylindrica Lemm. in 1954 (1), when it was Cyperaceae that small amounts of the element are Kylliniga brer ifolia Rottb. 57 Chlorosis 0.628 1.245 demonstrated specifically Amaranthaceae required for growth. Subsequently, sodium was shown to be Aniarauithus tricolor L. 40 Chlorosis 0.884 2.099 0.I essential for the angiosperm, A triplex vesicaria Heward ex cv. Early Splendour Benth (Bladder saltbush) (6), and more recently other species Chenopodiaceae been shown to to sodium re- Kochia clhildsii Hort. 21 Chlorosis 0.125 0.442 have respond (3, 16). Significant Portulacacea e sponses to sodium have been reported in barley (3) and to- Portulaca grandifiora 29 Clhlorosis, ne- 0.242 0.789 matoes (17), but the responses were small and no deficiency Hook (rose moss) crosis, and symptoms were observed. failure to Only certain species of Atriplex responded, decisively, to set flower i small quantities of sodium whereas other species of the same Names given in this table are those under which the seeds were received. genus showed no response (3). Subsequently, it was observed 2 Not significant. that the species of A triplex which required sodium had charac- teristics of plants with the C4 photosynthetic pathway whereas KH2PO4, 1,000; MgSO4, 1.000; (NH4)2HPO, 1,000: H3BO3, those not requiring sodium had features of the C3 photosyn- 46; MnSO,*7H20, 9.1; CuSO4 5H20, 0.31; ZnSO, 7H20, thetic pathway (9-11). 0.76; (NH4)5Mo7024A4H20, 0.1; NH4Cl, 350. Iron was sup- This communication gives results of experiments in which plied as ferric ammonium ethylenediaminetetraacetate at 90 six species from five families (Echinochloa utilis, Cynodon ,tmoles/liter in the basal culture solution. Less than 0.07 /.Leq/ dactylon, Kyllinga brevifolia, A maranthus tricolor, Kochia liter sodium was derived from the purified salts of the culture childsii, and Portulaca grandiflora) with features of C, plants solution as an impurity, and silica-distilled water contained less (7, 9) were shown to have a sodium requirement whereas Poa than 0.0087 lteq/liter, giving a total sodium concentration of pratensis, a species with C3 characteristics, showed no response approximately 0.08 ,ueq/liter. Culture vessels of 2-liter capac- to sodium. These results in conjunction with previously pub- ity were of sodium-free plastic material. Plants were grown in lished data are discussed to demonstrate a correlation between a naturally illuminated cabinet, slightly pressurized by a con- the essentiality of sodium and the possession of the Cl dicar- tinual supply of filtered air. This prevented the entry of dust boxylic photosynthetic pathway. particles and other atmospheric contaminants which could be 794 Plant Physiol. Vol. 49, 1972 REQUIREMENT FOR SODIUM AS A MICRONUTRIENT 795 Table II. Sodium Requirement in Relationi to C4 Pathway Characteristics Yield Plesints iNot _________ Signifi- Refer- C4 Pathway Probable o Species Receiving N 0.1 meq/ Difernce ne Caatrsis Reference Pathway Sodium liter _NaCI aditoadiin or Na2SO4 g dry wt/plant 7 Gramineae Hordeum vulgare L. cv. Pallidium None 0.77 0.99k 5 (barley) 3.82 4.01l (3) N,H (8) C3 Poa pratenisis L. (Kentucky blue grass) None 0.024 0.021 NS N C3 Echiniochloa utilis L. Ohwi et Yabuno Chlorosis and 0.404 0.713 0.1 K C4 (Japanese millet) necrosis Cynodon dactylon L. (Bermuda grass) Chlorosis 0.178 0.337 1 K,H14C4 (7) C4 Cyperaceae Kyllintga brevifolia Chlorosis 0.628 1.245 1 K C4 Amaranthaceae Amaranthus tricolor L. cv. Early Chlorosis and 0.884 2.099 0.1 K C4 Splendour necrosis Chenopodiaceae Chelnopodium capitatum L. Aschers None 12.25 14.37 NS (3) N (15) C3 Beta vulgaris L. (sugar beet) None 3.86 5.07 NS (3) N,H,L14C4 (8) C3 Atriplex nummularia Lindl. (oldman, Chlorosis 0.166 0.830 0.1 (3) K, L, H14C4, (11, 13, 14) C4 giant saltbush) L13C Atriplex paludosa R.Br. (marsh salt- Chlorosis and 0.215 2.789 5 (3) K C4 bush) necrosis Atriplex quinzii Fv.M. Chlorosis and 0.116 0.815 5 (3) K C4 necrosis Atriplex semibaccata R.Br. (berry salt- Chlorosis and 0.104 0.653 1 (3) K, L13C (13, 15) C4 bush) necrosis Atriplex inflata Fv.M. Chlorosis and 0.202 9.745 0.1 (3) K, H4C4 (11, 14) C4 necrosis Atriplex leptocarpa Fv.M. Chlorosis and 0.050 1.329 1 (3) K C4 necrosis Atriplex sponzgiosa Fv.M. (pop salt- Chlorosis and 0.570 12.472 0.1 (3) K, L, P. carb (10, 14) C4 bush) necrosis Atriplex semiluntalaris Aellen Chlorosis and 0.098 0.526 0.1 (3) Unknown necrosis Atriplex lindleyi Moq. Chlorosis and 0.093 0.560 5 (3) Unknown necrosis Atriplex vesicaria Heward ex Benth Chlorosis and 0.013 0.129 0.1 (2) K,H14C4, P. (11, 13) C4 (bladder saltbush) necrosis carb, L13C Atriplex hortenisis L. var. Atrosan- None 2.849 3.677 S (3) N,H,L14C4, (14, 15) C3 guineae (garden orache) H13C Atriplex angustifolia Sm. None 0.531 0.377 NS (3) Unknown ... Atriplex glabriuscula Edmonton None 24.804 24.100 NS (3) N,H,H13C (14) C3 Atriplex albicans Ait. None 8.818 11.688 9.237 9.401 NS (3) Unknown Kochia pyramidata Benth. None 35.390 33.990 NS (3) NIL14C4 (11) Ca Kochia childsii Hort. Chlorosis and 0.125 0.442 K,L,L13C (13, 14) C4 necrosis 1 Exomis axyrioides Fenzl ex Moq. None 0.864 0.925 NS (3) Unknown Halogeton glomeratus (Bieb) Meyer Smaller curved 0.285 0.800 (16) K (15) C3 leaves, tend- 1 ency to wilt- ing Cruciferae Brassica oleracea L. cv. "Savoy" None 14.244 17.766 NS (3) N (cabbage) C3 Leguminosae Trifolium repens L. cv. "Palestine" None 2.299 2.897 NS (3) N (white clover) C3 Solanaceae Lycopersicum esculentum Mill. cv. "Grosse Lisse" None 4.84 4.76 6.93 7.83 NS (3) N C3 cv. "Marglobe" None 13.76 15 .402 1 (17) N Ca 796 BROWNELL AND CROSSLAND Plant Physiol. Vol. 49, 1972 Table II-Continuiied Yield Lesions in Species Plants -Not Sinf- Refer- C4 Pathway Rfrne Probable Receiving N 0 e cance of ence Characteristics Pathway Sodium addition liter NaCl Difference or -Na2SO4 g dry wi/plant Compositae Lactuca sativa L. cv. "Great Lakes" None 3.871 6.113 (lettuce) 0.822 1 .228k NS (3) N C3 2.230 2.444J Aster tripolium L. None 0.282 0.458 NS (3)) Unknown Portulacaceae Portiulaca granidiflora Hook (rose Ilmoss) Chlorosis, no 0.242 0.789 1 K C4 flowers Leaf anatomy: N, normal (bifacial); K, Kranz. CO2 compensation: L, low; H, high. 'IC in C4 compounds (malate, aspartate): L"C4, low; H'4C4, high. Phosphoenol pyruvate carboxylase activity: P.carb, high. '3C discrimination: L"3C, low; H'3C, high. I Cultures contained 1 meq/liter NaCl. FIG. 1. Comparisons between plants of Echiniochloa uttilis (left), Amaranittlhs tricolor (center), and Kochia childsii (right) which received no addition (-Na) and 0.10 meq/liter sodium chloride (+Na). a source of sodium to the plants. Seeds were thoroughly washed (13). The known C, pathway characteristics for each species many times with distilled water before placing them in germi- are shown in Table II. No data relating to the C, pathway was nation cultures. The seedlings were transferred to their cul- available for certain species, including Atriplex setnilutnalaris, tures when sufficiently developed, and the sodium chloride Atriplex lindleyi, Atriplex angustifolia, Atriplex albicans, Ex- treatment was supplied to the appropriate cultures.
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