Nitrate Uptake by Manganese-Deficient Lemon Plants

The uptake of nitrate and water was followed in Eureka lemon (Citrus limon (L.) Burm. f.) plants grown in solution culture in a greenhouse under short (3–6 months) and long-term (22–24 months) Mn-nutrition stress. Uptake was determined from depletion in the nutrient solution. Under short-term stress, manganese-deficient plants absorbed 14.5% more nitrate and 3.4% more water than the control plants, on a weight basis. Under long-term stress there was a three-fold increase in nitrate and a two-fold increase in water uptake in the Mn-deficient plants. The intensive nitrate uptake under Mn-deficiency stress was more spectacularly demonstrated in plants which were exposed also to low nitrogen supply. The low-nitrogen Mn-deficient plants absorbed more nitrate, had less stunted growth and developed fewer visible symptoms of both N and Mn deficiencies than high-nitrogen Mn-deficient plants.     

Chromosomal polymorphisms in subterranean mole rats: origins and evolutionary significance

Two distinct chromosomal polymorphisms, among others, characterize the short (p) and long (qh) arms of chromosome number 1 of the Spalax ehrenbergi complex in Israel. We have studied the geographic distributions of these two polymorphisms in 60 animals belonging to four chromosomal species (2 n = 52, 54, 58 and 60). These comprised 15 populations: 12 from continuously distributed populations, two from semi-isolates, and one isolate in the northern Negev Desert.
Our results indicate that: (i) the two polymorphisms are widespread not only within populations and species but most strikingly between different bone marrow cells within an individual; this may reflect a diversity generating mechanism; (ii) the two polymorphisms of p and qh, primarily of p, are correlated with climatic factors of water availability and temperature, as well as geographical distances; these correlations are significantly above those expected by chance, (iii) The semi-isolates and the isolate display high levels of polymorphism in both p and qh, but particularly in qh, despite the small size population effects.
We suggest that at least the polymorphism in p is involved in the adaptive radiation of mole rats into diverse climatic regimes. Furthermore, we speculate that the forces generating chromosomal polymorphisms relevant to speciation (i.e. Robertsonian mutations), coupled with the evolutionary forces operating in small peripherally isolated populations, may be appropriate for the origin of new chromosomal species through peripatric speciation.     

Nitrogen Constituents in Manganese-Deficient Lemon Leaves

Total (Kjeldahl) nitrogen content of manganese-deficient lemon leaves was found to be about 40% higher (on a dry weight basis) than in the analog control leaves. The nitrate-N content rose in the deficient leaves from 18.2 mg/100 g to 95.6 mg/100 g dry weight. Among the N constituents studied, the free amino acids increased by about 42%, the protein amino acids by about 22%, and the total protein by more than 22%, all on a fresh weight basis. Acrylamide gel electrophoresis and ammonium sulfate fractionation showed qualitative differences in the various proteins. The increase of the protein fraction in the -Mn leaves may be attributed in part to the decrease of dry matter in these leaves (25.8% as compared to 30.7% in the control leaves) and in part to accumulation of apparent proteinous material in manganese-deficient tissues.     

Genetic parallelism of protein polymorphism in nature: ecological test of the neutral theory of molecular evolution

We have conducted an ecological test of protein polymorphism in 13 unrelated genera of plants, invertebrates and vertebrates, involving 21 species, 142 populations and 5474 individuals. Each was tested, on average, for 27 enzymatic gene loci. These species varied in population size and structure, life histories and biogeographical origins, but they largely share a geographically short (260 km) and ecologically stressful gradient of increasing aridity in Israel, both eastward and (mainly) southward. We found genetic parallelism across most taxa, and most loci. Observed average heterozygosity, H , and gene diversity, He , were positively and overall significantly correlated with rainfall variation. This result corroborates the environmental theory of genetic diversity, primarily the niche-width variation hypothesis in both space and time. Our results are inconsistent with the neutral theory of molecular evolution and suggest that natural selection appears to be an important differentiating evolutionary force at the protein level.     

Speciation and adaptive radiation of subterranean mole rats, Spalax ehrenbergi superspecies, in Jordan

The major initial mechanism of speciation in subterranean blind mole rats, Spalacidae, is chromosomal, primarily through Robertsonian rearrangements. Here we highlight another scenario of chromosomal rearrangement leading to ecological speciation and adaptive radiation apparently initiated by pericentric inversions and genie divergence to different ecologies in mole rats in Jordan. We analysed karyotype, allozyme, size and ecological diversity across the range of mole rats in Jordan from mesic Irbid in the north to xeric Wadi Musa (Petra region) in the south, a transect of 250 km. We examined mole rats for chromosome ( N =71), size ( N =76), and allozyme ( N =67) diversities, encoded by 32 loci, in 12 populations of the Spalax ehrenbetgi superspecies in Jordan. By a combination of chromosome morphology, genetic distance, body size and ecogeography, we identified four new putative biological species. All species (except two animals in Madaba) share 2 n = 60 but vary in chromosome morphology, caused by pericentric inversions and/or centromeric shifts. The 'north Moav' species is karyotypically polymorphic for 2 n (2 n = 60; including locally also two animals with 2 n = 62). The distribution of the four species is associated with ecogeographical different domains and climatic diversity. Genetic diversity indices were low, but like chromosome arms (NFa) were positively correlated with aridity stress. Discriminant analysis correctly classified 91% of the individuals into the four species utilizing combinatorially chromosome, allozyme and size diversities. It is hypothesized that mole rat evolution underground is intimately associated with climatic diversity stress above ground.     

Effect of Manganese Deficiency on Chloroplasts of Lemon Leaves

Manganese deficiency in chloroplasts of Eureka lemon leaves resulted in 23% and 40% increase of chloroplast nitrogen and protein, respectively, on a chlorophyll unit basis. Acrylamide gel electrophoresis carried out on extracts of these chloroplasts disclosed also qualitative differences between the normal and deficient leaves. Calculated on chloroplast N basis there is an increase of 17% in the chloroplast protein under Mn deficiency. This increase apparently indicates a more intense protein synthesis in the Mn deficient chloroplasts. Hill activity of the –Mn leaves was about one-third of the analog control leaves. Manganese infiltration into detached but intact leaves restored the activity in the –Mn leaves up to 70% of the control. Lemon leaves affected by other macro- and micro- nutrient deficiencies did not respond to the manganese infiltration; therefore, the use of this infiltration method is suggested for the evaluation of the manganese nutrition status of citrus and probably other higher plants.     

Karyotype and genetic evolution in speciation of subterranean mole rats of the genus Spalax in Turkey

Karyotype (2fl) and allozymc diversity at 37 gene loci were determined in 69 subterranean mole rats in Turkey belonging to the two superspecies: the ancestor Spalax leucodon ( n = 55; 20 populations) and the descendant S. ehrenbergi ( n = 14; lour populations). We identified remarkable variation of diploid chromosome numbers in the S. leucodon superspecies: 2 n = 38. 40. 50, 54, 60 and 62; and in the S. ehrenbergi superspecies: 2 n = 52, 56 and 58. Genetic diversity indices were low on average in both S. leucodon and S. ehrenbergi superspecies: Allele diversity, A = 1.081 and 1.074; polymorphism, P-5%= 0.077 and 0.068; heterozygosity, H = 0.038 and 0.027; and gene diversity, H, = 0.038 and 0.034, respectively. H ranged from 0 in mesic or semimesic regions to 0.088 in arid Anatolia. We consider the populations with different diploid chromosome numbers, 2 n , as good biological species. Karyotypic diversity may mark extensive ecological speciation. Nei's genetic distances, D (average 0.174, range 0.002 0.422) and ecogeographical criteria suggest that almost each population may represent a different biological species, but critical future testing is necessary to support this claim. Karyotypes and allozymes are nonrandomly distributed across Turkey, displaying remarkable correlations with climatic and biolir factors. Both In and H are significantly correlated with aridity stress (2 n /rainfall, r , =–0.74; P < 0.001), and in our region also with climatic unpredictability. These results support the niche-width genetic variation hypothesis in space and time. Climatic selection in Turkey appears to be a major architect of karyotype and genetic (allozymc) diversity and divergence in mole rat evolution, in both speciation and adaptation.     

Avoiding injury and surviving injury: two coexisting evolutionary strategies in lizards

Congenital morphological differences between injured and intact individuals in a population may reflect adaptations to avoid injury, to survive injury, or both. We explore the possible occurrence of such adaptations, analysing the relationship between tail state (original, O; regenerated, R) and morphology (20 scale characters) in 23 taxa of Lacertidae. In some taxa, such as Acanthodactylus opheodurus , morphologies of O and R lizards differed significantly. In these, usually O individuals were morphologically typical (modal), while R individuals were extreme; in others, the contrary occurred. Indices of pattern similarity detected fewer differences between O and R lizards than those based on absolute values. We developed unbiased estimates of classification rates of discriminant analysis. The order of inclusion of characters in the discriminant functions of males and females of the same species differed more than the order for the same sex in other species. Some morphological differences between O and R lizards seem adaptive or linked by pleiotropy to adaptive traits. Congenital morphological differences between O and R lizards are probably more frequent than detected because we show that age and geographical heterogeneity of our samples are confounding factors to O–R differences. R-morphologies might reflect microevolution in favour of paedotypic forms, possibly possessing greater regenerating capacities than relatively more peratypic forms.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 307–324.     

MtDNA polymorphisms: evolutionary significance in adaptation and speciation of subterranean mole rats

Patterns of mtDNA diversity in subterranean mole rats of the Spalax ehrenbergi superspecies (2n = 52, 54, 58 and 60) were previously studied in the main ranges of the four chromosomal species, and specifically in the 2n = 60 species and its peripheral steppe semi-isolates and desert isolates. In the present study we correlated mtDNA diversity indices, nucleon diversity, h , and nucleotide divergence, π, with physical (climatic), biotic (parasites) and biological (genetical, morphological, physiological and behavioural) factors, showing that mtDNA diversity is structured ecogeographically and biologically. The following significant correlations of mtDNA diversity were indicated with: (i) climatic heterogeneity and unpredictability; (ii) levels of ecto- and endoparasites; and (iii) biological diversities, primarily with physiological diversity associated with the energy budget. Small steppe semi-isolates and desert isolates harbour high levels of mtDNA haplotype diversity, some novel, which may be a prerequisite for future speciation events. We conclude that the ecogeographical and biological correlates, as well as the maintenance of mtDNA polymorphisms in small isolated populations, strongly suggest that mtDNA diversity is not neutral. Diversifying natural selection appears to be an important differentiating factor of mtDNA diversity in the twin evolutionary processes of adaptive radiation and active speciation. We suggest critical experiments to substantiate our conclusions and highlight the contribution of mtDNA diversity to fitness, i.e. to the biological function of mtDNA diversity in the evolutionary process.