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Heterogeneity in Carbon Isotope Discrimination in Leaves, Stalks and Spikes of Ten Annual Wild Triticeae Species

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Heterogeneity in Carbon Isotope Discrimination in Leaves, Stalks and Spikes of Ten Annual Wild Triticeae Species Czech J. Genet. Plant Breed., 41, 2005 (Special Issue) Heterogeneity in Carbon Isotope Discrimination in Leaves, Stalks and Spikes of Ten Annual Wild Triticeae Species J. Z������� and V. H������ Research Institute of Crop Production, 161 06 Praha-Ruzyně, Czech Republic e-mail: [email protected], [email protected] Abstract: The wild Triticeae species are confined mostly to arid regions and thus, they are suitable material for study of their adaptation to dry conditions. Species from semidesert localities of Mediterranean, Balkans and Near East (genera Eremopyrum, Heteranthelium, Taeniatherum, Dasypyrum, and Aegilops) and wild wheats (Triticum) from steppe localities were studied. The studied species can be divided into two groups on the basis of their carbon isotope discrimination (Δ). The first group, which had high Δ, consists of species (Ta. crinitum, A. tauschii, H. piliferum, E. orientale, A. markgrafii, A. comosa) collected in the driest localities such as semidesert sites. The second group from steppe localities, which consists of D. villosum, T. urartu, T. boeoticum, and T. monococcum, had relatively low Δ. We conclude that wild species originally grown in dry regions discriminate more carbon isotope as they are adapted to the dry condition by stomata opening and/or Rubisco activity. We examined from high to low carbon isotope discrimination values of the flag leaves, stalks, and spikes. Keywords: carbon isotope discrimination; delta; wild Triticeae; heterogeneity; drought adaptation Wild wheat relatives, members of the Triticeae ing method for evaluation transpiration efficiency tribe, could represent a valuable source of genetic (TE). Δ is a measure of the ratio of stable carbon variation for improvement of abiotic stress tolerance isotopes (13C/12C) in plant dry ma�er compared in cultivated wheat (Triticum aestivum L.). A better with the same ratio in the atmosphere. 13 knowledge of the adaptive strategies developed Discrimination against CO2 is closely negatively by these species is needed. Species of Aegilops correlated with the transpiration efficiency inte- (goatgrass) are well adapted to dry environments. grated over the life of the sampled plant material Aegilops tauschii is of particular interest, since gene (F������� & R������� 1984; C����� et al. 1990; transfer from this drought-adapted species to bread C����� et al. 1992). Carbon isotope discrimination wheat is relatively easy (V������ et al. 2004). has been proposed as a useful indicator of TE in Isotopic differences in diverse elements make it wheat (F������� & R������� 1984; E����� et al. possible to demonstrate that fractionation events 1991) and in numerous species including other occur in nature. Fractionation processes are ther- cereals (C����� et al. 1993; M���� et al. 1996). The modynamic or kinetic reactions that produce dif- negative correlation of Δ and water use efficiency ferent abundances in reactants and products. Two (WUE, measured either as net photosynthesis/ 13 important processes in which CO is discriminated transpiration or plant biomass produced/water 2 are diffusion and carboxylation. Diffusion fractiona- transpired) led to Δ being proposed as selection tion is dependent on molecular mass. Diffusion of criteria for WUE (F������� & R������� 1984). It heavy atoms or molecules is slower than diffusion of is determined by the intercellular partial pressure lighter ones. Carboxylation by RUBISCO discrimi- of CO2, which is largely regulated by variation in nates strongly against 13CO typically in C plants. stomatal aperture (F������� et al. 1989). Under 2 3 Carbon isotope discrimination (Δ) is a very promis- drought stress, Δ can be considered as a good 212 Proc. 5th International Triticeae Symposium, Prague, June 6–10, 2005 Czech J. Genet. Plant Breed., 41, 2005 (Special Issue) predictor of stomatal conductance (C����� et al. tope discrimination and the total carbon content 1990). Carbon isotope discrimination positively in plant parts. correlates with yield and growth cycle duration (A���� et al. 1997) and negatively correlates with MATERIAL AND METHODS leaf temperature (A������ 1993). The strong corre- lation between carbon isotope discrimination and Material. Mature plants of wild Eremopyrum, yield in durum wheat grown in the Mediterranean Heteranthelium, Taeniatherum, Dasypyrum, Triticum conditions has been confirmed (H���� et al. 2000). and Aegilops species were collected at the end of It is easier to measure carbon isotope discrimina- vegetation period from semidesert and steppe tion than transpiration efficiency. Measuring of localities of the Mediterranean, Balkans and Near carbon isotope ratio in dry matter of a plant organ East from steppe localities (Table 1). The localities provides a measure of the discrimination of carbon are characterized by a dry summer season with isotope of this organ and carbon isotope ratio of high temperature and high irradiance. matter coming from other plant organs. Methods. Collected plants were dried until they The main objectives of the present study were (i) reached a constant weight. Twenty flag leaf blades, to examine the variation in plant carbon isotope twenty 10 cm sections from the upper part of the discrimination as a tool which represents water stalk, and twenty spikes per genotype were ran- use efficiency among accessions of the wild Tri- domly sampled from fully mature plants. The leaf, ticeae collected in their native habitats where they stalk and spike samples were separately grounded are well acclimated (ii) to analyze relationships in to a fine powder in a metal ball mill. Carbon isotope carbon isotope discrimination of different wild ratios were measured to 0.10‰ by isotope ratio mass Triticeae organs in generative stage important for spectrometer (IRMS). The total carbon content and reproduction at the end of plant ontogenesis and carbon isotope composition of leaves stalks and (iii) to evaluate relationship between carbon iso- spikes were determined by an Elemental Analyzer Table 1. Species used for study, their geographic site and ecological characteristic Accession Abbrevia- Collector/ Latin Name Geographic Site Ecology No. tion donor C2100705 Aegilops comosa SIBTH. et SM. A. com NW Korfu on Isle of saline grassland Gatersleben ssp. heldreichii (BOISS.) EIG Korfu, Near Potamos River Aegilops markgrafii (GREUTER) Hasan Dag, nr. Hamrun, bazalt, lava pall, C2100428 A. mar VH HAMMER var. markgrafii Nigde, Turkey 1460 m TA2457 Aegilops tauschii COSS. A. tau Turkey dry steppe Kansas Dasypyrum villosum (L.) Sopot, 7 km N of C2300014 D. vil dry steppe, 415 m VH CANDARGY Kumanovo, Macedonia Eremopyrum orientale (L.) JAUB. Dzrvez, 2 km E of Erevan semidesert steppe, C3400006 E. ori VH et SPACH to Garni, Armenia 1000 m C3700001 Heteranthelium piliferum H. pil Iran semidesert Logan (BANKS et SOLANDER) HOCHST. Triticum boeoticum BOISS. var. Izgrev – Pcela, Rodopi dry steppe, 680 m VH C0104087 T. boe1 symbolense (Flaksb.) A. Filat. Mts., Bulgaria C0106065 Triticum boeoticum BOISS. T. boe2 Bayindr, Konya, Turkey dry steppe, 1195 m VH Taeniatherum crinitum Ni�aular nr. Dzebraili, semidesert steppe, C3300006 Ta. cri VH (SCHREBER) NEVSKI Georgia 1000 m C0106421 Triticum monococcum L. var. T. mon Transylvania, Romania old land race A. Szabo clusianum LR. Galara Kiss Triticum urartu THUM. ex dry steppe to C0106076 T. ura Sanli Urfa, Turkey ICARDA GANDIL. semidesert, 620 m VH = Vojtech Holubec Proc. 5th International Triticeae Symposium, Prague, June 6–10, 2005 213 Czech J. Genet. Plant Breed., (EuroVector 3028/HT) coupled to an Isotope Ratio Mass Spectrometer (IRMS Isoprime). International standards were used so all delta values are inter- comparable. We use delta notation ( carbon isotope ratio conveniently: δ 13 C = ([( delta values are expressed in parts 41, 2005 (Special Issue) (‰ 13 number. C/ ) for plant material12 C) on average.The standard Fractionation error of is determination also known as was carbon 0.10 isotope discrimination ( sample ples was obtained according to the formula:/( 13 C/ Δ ≈ ( 12 C) Δ δ δ ≈ a – ∂ standard ) to express 13 ( δ δ F�������source C is always a negative p )/(1 + ] plant carbon isotope composition, respectively. The – carbon isotope composition – of air was 1 ) Statistica 6.1 (StatSoft) software was used for of C δ δ Δ 1000 statistical evaluation of the data. product p ), ). The sample was analyzed at least et al. 2 times. Direct transpi “ (1) ration efficiency3 plants was is reportednot measured approximately in this study. per mil 1989 Δ value of the sam overallTaeniatherum crinitum ), ” RESULTS AND DISCUSSION where a and p refer to air and cum monococcum Dasypyrum ‰ carbon isotope discrimination.carbon isotope Interestingly, discrimination and crinitum - is characteristic for species with low water use efficiency. The and all studied had low carbon discrimination, corresponding to 21 (2) had values of more the thanlowest (Figure 1). In general, the high WUE. – (3) exhibited the highest 13 C (‰) 8.3 positionFor mostof the of leaves the species was lowstudied, in comparison the carbon comto � – ‰ 25 rest of the plant 20 ‰ , δ carbon composition of 48–50% in their stalks and . Each 13 Triticum Triticum C spikes. were exceptional in this respect, having values of only - Aegilops comosa spp. on the other hand, had the lowest 19 composition of the leaves of with the values37% obtainedand 30%, for respectively. its stalks and spikes Δ Triti (Figure 3) (Figure 2). 20 Carbon Isotope Discrimination ‰ - 18 , which Ta. and Figure 1 Average carbon isotope discrimination of
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