Central Arid Zone Research Institute
governmentJodhpur, India
Research output, citation impact, and the most-cited recent papers from Central Arid Zone Research Institute (India). Aggregated across the NobleBlocks index of 300M+ scholarly works.
Top-cited papers from Central Arid Zone Research Institute
Lead (Pb) toxicity has been a subject of interest for environmental scientists due to its toxic effect on plants, animals, and humans. An increase in several Pb related industrial activities and use of Pb containing products such as agrochemicals, oil and paint, mining, etc. can lead to Pb contamination in the environment and thereby, can enter the food chain. Being one of the most toxic heavy metals, Pb ingestion via the food chain has proven to be a potential health hazard for plants and humans. The current review aims to summarize the research updates on Pb toxicity and its effects on plants, soil, and human health. Relevant literature from the past 20 years encompassing comprehensive details on Pb toxicity has been considered with key issues such as i) Pb bioavailability in soil, ii) Pb biomagnification, and iii) Pb- remediation, which has been addressed in detail through physical, chemical, and biological lenses. In the review, among different Pb-remediation approaches, we have highlighted certain advanced approaches such as microbial assisted phytoremediation which could possibly minimize the Pb load from the resources in a sustainable manner and would be a viable option to ensure a safe food production system.
Biological synthesis of ZnO nanoparticle is a new approach for environmentally benign protocol in context to green nanotechnology. In present investigation, ZnO nanoparticles were synthesized from ZnNO3 using extracellular secretions of Aspergillus fumigatus TFR-8 (NCBI GenBank Accession No. JQ675291) and effect of these biologically transformed ZnO nanoparticles was studied on clusterbean (Cyamopsis tetragonoloba L.) to enhance native phosphorous-mobilizing enzymes and nanoinduced gum production. Valid characterization techniques were employed for confirmation of size, shape, surface structure, crystalline nature, and elemental proportion. Results indicate that synthesized nanoparticle size ranging between 1.2 and 6.8 nm at least in one dimension with oblate spherical and hexagonal in structure. The samples contained 98 % atom of Zn element. The characterized ZnO nanoparticles were foliar sprayed at 10 ppm concentration on leaf of 14-day-old clusterbean plants. A significant improvement in plant biomass (27.1 %), shoot length (31.5 %), root length (66.3 %), root area (73.5 %), chlorophyll content (276.2 %), total soluble leaf protein (27.1 %), rhizospheric microbial population (11–14 %), acid phosphatase (73.5 %), alkaline phosphatase (48.7 %), and phytase (72.4 %) activity in clusterbean rhizosphere was observed over control in 6-week-old plants due to application of nanoZnO. The gum content in clusterbean seeds improved by 7.5 % after maturity which indicates ZnO in nano form may contribute more in industrial and medical applications besides agricultural sector. A possible hypothesis of mechanism for ZnO nanoparticle biosynthesis has also been made.
Abstract Pearl millet [ Cenchrus americanus (L.) Morrone] is a staple food for more than 90 million farmers in arid and semi-arid regions of sub-Saharan Africa, India and South Asia. We report the ∼1.79 Gb draft whole genome sequence of reference genotype Tift 23D 2 B 1 -P1-P5, which contains an estimated 38,579 genes. We highlight the substantial enrichment for wax biosynthesis genes, which may contribute to heat and drought tolerance in this crop. We resequenced and analyzed 994 pearl millet lines, enabling insights into population structure, genetic diversity and domestication. We use these resequencing data to establish marker trait associations for genomic selection, to define heterotic pools, and to predict hybrid performance. We believe that these resources should empower researchers and breeders to improve this important staple crop.
The present study demonstrates an eco-friendly and low cost protocol for synthesis of silver nanoparticles using the cell-free filtrate of Aspergillus flavus NJP08 when supplied with aqueous silver (Ag+) ions. Identification of the fungal isolate was based on nuclear ribosomal DNA internal transcribed spacer (ITS) identities. Transmission electron microscopy (TEM) and energy dispersive spectroscopy (EDS) revealed the formation of spherical metallic silver nanoparticles. The average particle size calculated using Dynamic Light Scattering measurements (DLS) was found to be 17±5.9 nm. UV-Visible and Fourier transform infrared (FTIR) spectroscopy confirmed the presence of extracellular proteins. SDS-PAGE profiles of the extracellular proteins showed the presence of two intense bands of 32 and 35 kDa, responsible for the synthesis and stability of silver nanoparticles, respectively. A probable mechanism behind the biosynthesis is discussed, which leads to the possibility of using the present protocol in future "nano-factories".
KEY MESSAGE: Analysis of phenotypic data for 20 drought tolerance traits in 1-7 seasons at 1-5 locations together with genetic mapping data for two mapping populations provided 9 QTL clusters of which one present on CaLG04 has a high potential to enhance drought tolerance in chickpea improvement. Chickpea (Cicer arietinum L.) is the second most important grain legume cultivated by resource poor farmers in the arid and semi-arid regions of the world. Drought is one of the major constraints leading up to 50% production losses in chickpea. In order to dissect the complex nature of drought tolerance and to use genomics tools for enhancing yield of chickpea under drought conditions, two mapping populations-ICCRIL03 (ICC 4958 × ICC 1882) and ICCRIL04 (ICC 283 × ICC 8261) segregating for drought tolerance-related root traits were phenotyped for a total of 20 drought component traits in 1-7 seasons at 1-5 locations in India. Individual genetic maps comprising 241 loci and 168 loci for ICCRIL03 and ICCRIL04, respectively, and a consensus genetic map comprising 352 loci were constructed ( http://cmap.icrisat.ac.in/cmap/sm/cp/varshney/). Analysis of extensive genotypic and precise phenotypic data revealed 45 robust main-effect QTLs (M-QTLs) explaining up to 58.20% phenotypic variation and 973 epistatic QTLs (E-QTLs) explaining up to 92.19% phenotypic variation for several target traits. Nine QTL clusters containing QTLs for several drought tolerance traits have been identified that can be targeted for molecular breeding. Among these clusters, one cluster harboring 48% robust M-QTLs for 12 traits and explaining about 58.20% phenotypic variation present on CaLG04 has been referred as "QTL-hotspot". This genomic region contains seven SSR markers (ICCM0249, NCPGR127, TAA170, NCPGR21, TR11, GA24 and STMS11). Introgression of this region into elite cultivars is expected to enhance drought tolerance in chickpea.
TiO2 nanoparticle (NPs) biosynthesis is a low cost, ecofriendly approach developed using the fungi Aspergillus flavus TFR 7. To determine whether TiO2 NPs is suitable for nutrient, we conducted a two part study; biosynthesis of TiO2 NP and evaluates their influence on mung bean. The characterized TiO2 NPs were foliar sprayed at 10 mgL−1 concentration on the leaves of 14 days old mung bean plants. A significant improvement was observed in shoot length (17.02%), root length (49.6%), root area (43%), root nodule (67.5%), chlorophyll content (46.4%) and total soluble leaf protein (94%) as a result of TiO2 NPs application. In the rhizosphere microbial population increased by 21.4–48.1% and activity of acid phosphatase (67.3%), alkaline phosphatase (72%), phytase (64%) and dehydrogenase (108.7%) enzyme was observed over control in six weeks old plants owing to application of TiO2 NPs. A possible mechanism has also been hypothesized for TiO2 NPs biosynthesis.
Ten days old seedlings of chickpea (Cicer arietinum L var. HC-1) were foliar sprayed with 1.5 or 10 ppm aqueous solution of zinc oxide (ZnO) nanoparticles and effects were compared with corresponding concentration of zinc sulphate and ZnO of normal size. Maximum promontory response with respect to shoot dry weight was observed in seedlings treated with 1.5 ppm ZnO nanoparticles while at 10 ppm the nanoparticles exerted adverse effects on root growth. However, overall biomass accumulation improved in the ZnO nanoparticle treated seedlings. This response may be attributed to low reactive oxygen species (ROS) levels which resulted in less lipid peroxidation as evident from lower malondialdehyde (MDA) content. This was associated with lower activity of prominent antioxidant enzymes, superoxide dismutase (SOD), and peroxidase ZnO nanoparticle treated seedling compared to control. The study indicates importance in precise application of zinc, more so in deficient system, where plant response varies with concentration and is important in understanding the mechanism of action of specific nanomaterials.
Phosphorus (P) is a limiting factor to plant growth and productivity in almost half of the world's arable soil, and its uptake in plants is often constrained because of its low solubility in the soil. To avoid repeated and large quantity application of rock phosphate as a P fertilizer and enhance the availability of native P acquisition by the plant root surface, in this study a biosynthesized ZnO nanoparticle was used. Zn acts as a cofactor for P-solubilizing enzymes such as phosphatase and phytase, and nano ZnO increased their activity between 84 and 108%. The level of resultant P uptake in mung bean increased by 10.8%. In addition, biosynthesized ZnO also improves plant phenology such as stem height, root volume, and biochemical indicators such as leaf protein and chlorophyll contents. In the rhizosphere, increased chlorophyll content and root volume attract microbial populations that maintain soil biological health. ICP-MS results showed ZnO nanoparticles were distributed in all plant parts, including seeds. However, the concentration of Zn was within the limit of the dietary recommendation. To the best of our knowledge, this is the first holistic study focusing on native P mobilization using ZnO nanoparticles in the life cycle of mung bean plants.
Climate models project that many terrestrial ecosystems will become drier over the course of this century, leading to a drastic increase in the global extent of arid soils. In order to decrease the effects of climate change on global food security, it is crucial to understand the arid environment and the constraints associated with arid soils. Although the effects of aridity on aboveground organisms have been studied extensively, our understanding of how it affects soil processes and nutrient cycling is lacking. One of the primary agricultural constraints, particularly in arid locations, is water scarcity, due to which arid soils are characterized by sparse vegetation cover, low soil organic carbon, poor soil structure, reduced soil biodiversity, and a high rate of soil erosion via wind. Increased aridity will limit the availability of essential plant nutrients and crop growth, and subsequently pose serious threats to key ecological processes and services. The increasing rate of soil salinization is another major environmental hazard that further limits the agricultural potential of arid soils. These soil constraints can be ameliorated and the crop yields increased through case-specific optimization of irrigation and drainage management, enhancing the native beneficial soil microbes, and combinations of soil amendments, conditioners, and residue management. This review explores technologies to ameliorate soil constraints and increase yields to maintain crop output in arid soils.
Sustainable enhancement in food production from less available arable land must encompass a balanced use of inorganic, organic, and biofertilizer sources of plant nutrients to augment and maintain soil fertility and productivity. The varied responses of microbial inoculants across fields and crops, however, have formed a major bottleneck that hinders its widespread adoption. This necessitates an intricate analysis of the inter-relationships between soil microbial communities and their impact on host plant productivity. The concept of “biased rhizosphere,” which evolved from the interactions among different components of the rhizosphere including plant roots and soil microflora, strives to garner a better understanding of the complex rhizospheric intercommunications. Moreover, knowledge on rhizosphere microbiome is essential for developing strategies for shaping the rhizosphere to benefit the plants. With the advent of molecular and “omics” tools, a better understanding of the plant-microbe association could be acquired which could play a crucial role in drafting the future “biofertilizers.” The present review, therefore aims to (a) to introduce the concepts of rhizosphere hotspots and microbiomes and (b) to detail out the methodologies for creating biased rhizospheres for plant-mediated selection of beneficial microorganisms and their roles in improving plant performance.
Drought is one of the most prevalent limiting factors causing considerable losses in crop productivity, inflicting economic as well as nutritional insecurity. One of the greatest challenges faced by the scientific community in the next few years is to minimize the yield losses caused by drought. Drought resistance is a complex quantitative trait controlled by many genes. Thus, introgression of drought resistance traits into high yielding genotypes has been a challenge to plant breeders. Vegetable grafting using rootstocks has emerged as a rapid tool in tailoring plants to better adapt to suboptimal growing conditions. This has induced changes in shoot physiology. Grafting applications have expanded mainly in Solanaceous crops and cucurbits, which are commonly grown in arid and semi-arid areas characterized by long drought periods. The current review gives an overview of the recent scientific literature on root-shoot interaction and rootstock-driven alteration of growth, yield, and fruit quality in grafted vegetable plants under drought stress. Further, we elucidate the drought resistance mechanisms of grafted vegetables at the morpho-physiological, biochemical, and molecular levels.
Nanotechnology is the study of particle sizes between 1 and 100 nanometers at least at one dimension. Particle size reduced to nanometer length scale exhibit more surface area to volume size ratio and showing unusual properties makes them enable for systematic applications in engineering, biomedical, agricultural and allied sectors. Nanomaterial can create from bottom up or top down approaches using physical, chemical and biological mode of synthesis. Key words: Nanotechnology, nanomaterial, nanobiotechnology, nanotech-applications.
Water quality evaluation is critically important for the protection and sustainable management of groundwater resources, which are variably vulnerable to ever-increasing human-induced physical and chemical pressures (e.g., overexploitation and pollution of aquifers) and to climate change/variability. Preceding studies have applied a variety of tools and techniques, ranging from conventional to modern, for characterization of the groundwater quality worldwide. Recently, geographic information system (GIS) technology has been successfully integrated with the advanced statistical/geostatistical methods, providing improved interpretation capabilities for the assessment of the water quality over different spatial scales. This review intends to examine the current standing of the GIS-integrated statistical/geostatistical methods applied in hydrogeochemical studies. In this paper, we focus on applications of the time series modeling, multivariate statistical/geostatistical analyses, and artificial intelligence techniques used for groundwater quality evaluation and aquifer vulnerability assessment. In addition, we provide an overview of salient groundwater quality indices developed over the years and employed for the assessment of groundwater quality across the globe. Then, limitations and research gaps of the past studies are outlined and perspectives of the future research needs are discussed. It is revealed that comprehensive applications of the GIS-integrated advanced statistical methods are generally rare in groundwater quality evaluations. One of the major challenges in future research will be implementing procedures of statistical methods in GIS software to enhance analysis capabilities for both spatial and temporal data (multiple sites/stations and time frames) in a simultaneous manner.
The bacterial genus Exiguobacterium accommodates many versatile species isolated from diverse environments. Exiguobacterium was described as a genus approximately three decades ago, and now, 17 species, growing over a broad range of temperatures and pH, have been recognized. Various isolates from different niches have been explored for biotechnological and industrial purposes, including enzyme production, bioremediation and degradation of toxic substances released into the environment. Some isolates possess plant growth promoting capabilities, and they are currently being explored for increasing agricultural production. The genome sequences of various strains of this genus have shown the presence of many genes encoding products of importance to agriculture and the environment. In addition, many strains possess stress-responsive genes helping them to colonize and thrive in diverse ecological niches. This review provides a broad view of the versatile genus Exiguobacterium and its potential for applications in agriculture, the environment and industry, as well as the underlying genomic determinants that drive its diversity and adaptability to various extreme environments.
Of late, intensive farming for higher food production is often associated with many negative implications for soil systems, such as decline of soil organic matter (SOM), increase in risks of soil erosion by wind and/or water, decline in soil biological diversity, increase in degradation of soil physical quality, lower nutrient-use efficiency, high risks of groundwater pollution, falling water tables, increasing salinization and waterlogging, in-field burning of crop residues, pollution of air and emission of greenhouse gases (GHG), leading to global warming, and decline in factor productivity. These negative implications necessitate an objective review of strategies to develop sustainable management practices, which could not only sustain soil health and ensure food security, but also enhance carbon sequestration, decrease GHG emissions, and offer clean and better ecosystem services. Conservation agriculture (CA), that includes reduced or no-till practices along with crop residue retention and mixed crop rotations, offers multiple benefits. Adoption of a system-based CA conserves water, improves and creates more efficient use of natural resources through the integrated management of available soil nutrients, water, and biological resources, and enhances use efficiency of external inputs. Due to apparent benefits of CA, it is increasingly being adopted and now covers about 180 million hectares (Mha) worldwide. However, in South Asia its spread is low (<5 Mha), mostly concentrated in the Indo-Gangetic Plains (IGP). In this region, one of the serious issues is “residue burning” with severe environmental impacts. A huge amount of crop residue left over after the combine harvest of rice has forced farmers to practice widespread residue burning (∼140 M tonnes) to cope with excessive stubble and also for timely planting/sowing of succeeding crops.In rice-wheat cropping systems, which cover more than 10 Mha in the IGP, CA practices are relatively more accepted by farmers. In these systems, any delay in sowing leads to yield penalty of 1–1.5% per day after the optimum sowing date of wheat. The strong adoption of CA practices in IGP is mainly to overcome delayed sowing due to the field preparation and control of weeds, timely planting, and also escape from terminal heat during the grain-filling stage. Major challenges to CA adoption in South Asia are small land holdings (<1 ha), low technological reach to farmers, nonavailability of suitable farm implements for small farm holders, and the staunch conventional farming mind-set. South Asia region consists of many countries of diverse agro-ecologies with contrasting farming systems and management. This region, recently known for rapid economic growth and increasing population, necessitates higher food production and also hot-spots for adoption of CA technologies. Therefore, in this review critically explores the possibility, extent of area, prospects, challenges, and benefits of CA in South Asia.HIGHLIGHTSConservation agriculture (CA), consisting of reduced or no-tillage and crop residue retention, is a self–sustainable system which offers an alternative to crop residue burning.The CA approach improves soil health by increasing soil organic carbon (SOC) and aggregation and also conserves soil, water and energy than conventional farming systems.South Asian countries are suitable for adoption of CA practices but the area under CA remains low (<5 Mha) as compared to the global area (180 Mha).Adoption of CA in South Asia has skewed distribution, mainly in Indo-Gangetic Plains (IGP) in India, Pakistan, Nepal and Bangladesh in South Asia.Development of herbicide-resistant weed species and weed shift by continuous application of herbicides are the major challenges in adoption of CA.The traditional-farming mind-set, socio-economic conditions, small farm-holdings, weed and residue management, and non-availability of suitable machinery are key constraints to the low adoption of CA practices in South Asia.
Abstract. In this paper, we present and analyze a novel global database of soil infiltration measurements, the Soil Water Infiltration Global (SWIG) database. In total, 5023 infiltration curves were collected across all continents in the SWIG database. These data were either provided and quality checked by the scientists who performed the experiments or they were digitized from published articles. Data from 54 different countries were included in the database with major contributions from Iran, China, and the USA. In addition to its extensive geographical coverage, the collected infiltration curves cover research from 1976 to late 2017. Basic information on measurement location and method, soil properties, and land use was gathered along with the infiltration data, making the database valuable for the development of pedotransfer functions (PTFs) for estimating soil hydraulic properties, for the evaluation of infiltration measurement methods, and for developing and validating infiltration models. Soil textural information (clay, silt, and sand content) is available for 3842 out of 5023 infiltration measurements (∼ 76%) covering nearly all soil USDA textural classes except for the sandy clay and silt classes. Information on land use is available for 76 % of the experimental sites with agricultural land use as the dominant type (∼ 40%). We are convinced that the SWIG database will allow for a better parameterization of the infiltration process in land surface models and for testing infiltration models. All collected data and related soil characteristics are provided online in *.xlsx and *.csv formats for reference, and we add a disclaimer that the database is for public domain use only and can be copied freely by referencing it. Supplementary data are available at https://doi.org/10.1594/PANGAEA.885492 (Rahmati et al., 2018). Data quality assessment is strongly advised prior to any use of this database. Finally, we would like to encourage scientists to extend and update the SWIG database by uploading new data to it.
In the present study, zinc (Zn), magnesium (Mg) and titanium (Ti) nanoparticles synthesized using fungus by employing various precursor salts of sulfate salts, nitrate salts, chloride salts and oxide salts. To access the nanoparticle production potential, over a hundreds of fungi were isolated from the soil and tested with precursor salts of the Zn, Mg and Ti. Out of which, only 14 fungal isolates were identified, having potential to reduce metal salt into metal nanoparticles. Upon molecular identification, six were identified as Aspergillus flavus, two each as Aspergillus terreus and Aspergillus tubingensis and one each as Aspergillus niger, Rhizoctonia bataticola, Aspergillus fumigatus, and Aspergillus oryzae. Factors responsible for more production of monodispersed Zn, Mg and Ti nanoparticles were optimized. It was concluded that 0.01 mM precursor salt concentration, 72 h of incubation at pH 5.5 and temperature 28 °C resulted smaller nanoparticles obtained. The biosynthesized functional Zn and Ti nanoparticles can be stored up to 90 days and Mg nanoparticles up to 105 days in its nanoform. Bio-transformed products were analyzed using valid characterization technique i.e. dynamic light scattering, transmission electron microscopy, atomic force microscopy, energy dispersive X-ray spectroscopy to confirm size, shape, surface morphology and elemental composition. It was found that the average size of developed nano Zn was 8.2 nm, with surface charge of −5.70 mV and 98 % particles were of Zn metal only. Similarly, the average size of Mg nanoparticles was 6.4 nm with surface charge of −6.66 and 97.4 % Mg metal yield, whereas, Ti nanoparticles size were found in the ranges between 1.5 and 30 nm with surface charge of −6.25 mV and 98.6 % Ti metal yield.
A study was conducted to demonstrate the comparative efficiency of acid phosphatase generated by plants or fungi towards the hydrolysis of different organic P compounds present in soil. The results revealed that acid phosphatases were most efficient in the hydrolysis of glycerophosphate followed by lecithin and phytin. The P release increased with increase in enzyme concentration. Acid phosphatase generated from fungal sources showed three times greater efficiency in the hydrolysis of phytin, two times greater efficiency in hydrolysis of lecithin than plant phosphatase. Both sources were at par in hydrolyzing glycerophosphate. The results suggest that acid phosphatase generated from plant and fungal sources is different and microbial acid phosphatase to be more efficient than that from plant sources. Wirkungsvergleich saurer Phosphatasen pflanzlichen und pilzlichen Ursprungs Die sauren Phosphatasen aus Sorghum bicolor, Vigna unguiculata und Phaseolus radiata, sowie aus Aspergillus niger, A. terreus und A. rugulosus wurden bezüglich der Hydrolyse von Phytin, Lecithin und Glycerinphosphat verglichen. Die P-Freisetzung war linear abhängig von der Enzymkonzentration. Am leichtesten wurde Glycerinphosphat abgebaut, gefolgt von Lecithin und Phytin. Die Phosphatase aus Aspergillus schloss Phytin dreifach und Lecithin zweifach effizienter auf als die pflanzlichen Phosphatasen, während bei Glycerinphosphat keine Wirkungsunterschiede beobachtet wurden.
MicroRNAs (miRNAs) are important for plant development and stress responses. However, factors regulating miRNA metabolism are not completely understood. SICKLE (SIC), a proline-rich protein critical for development and abiotic stress tolerance of Arabidopsis, was identified in this study. Loss-of-function sic-1 mutant plants exhibited a serrated, sickle-like leaf margin, reduced height, delayed flowering, and abnormal inflorescence phyllotaxy, which are common characteristics of mutants involved in miRNA biogenesis. The sic-1 mutant plants accumulated lower levels of a subset of miRNAs and transacting siRNAs but higher levels of corresponding primary miRNAs than the WT. The SIC protein colocalizes with the miRNA biogenesis component HYL1 in distinct subnuclear bodies. sic-1 mutant plants also accumulated higher levels of introns from hundreds of loci. In addition, sic-1 mutant plants are hypersensitive to chilling and salt stresses. These results suggest that SIC is a unique factor required for the biogenesis of some miRNAs and degradation of some spliced introns and important for plant development and abiotic stress responses.
Late Pleistocene terrestrial climate records in India may be preserved in oxygen and carbon stable isotopes in pedogenic calcrete. Petrography shows that calcrete nodules in Quaternary sediments of the Thar Desert in Rajasthan are pedogenic, with little evidence for postpedogenic alteration. The calcrete occurs in four laterally persistent and one nonpersistent eolian units, separated by colluvial gravel. Thermoluminescence and infrared- and green-light-stimulated luminescence of host quartz and feldspar grains gave age brackets for persistent eolian units I–IV of ca. 70,000–60,000, ca. 60,000–55,000, ca. 55,000–43,000, and ca. 43,000–∼25,000 yr, respectively. The youngest eolian unit (V) is <10,000 yr old and contains no calcrete. Stable oxygen isotope compositions of calcretes in most of eolian unit I, in the upper part of eolian unit IV, and in the nonpersistent eolian unit, range between −4.6 and −2.1‰ PDB. These values, up to 4.4‰ greater than values from eolian units II and III, are interpreted as representing nonmonsoonal 18 O-enriched “normal continental” waters during climatic phases when the monsoon weakened or failed. Conversely, 25,000–60,000-yr-old calcretes (eolian units II and III) probably formed under monsoonal conditions. The two periods of weakened monsoon are consistent with other paleoclimatic data from India and may represent widespread aridity on the Indian subcontinent during isotope stages 2 and 4. The total variation in δ 13 C is 1.7‰ (0.0–1.7‰), and δ 13 C covaries positively and linearly with δ 18 O. δ 13 C values are highest when δ 18 O values indicate the most arid climatic conditions. This is best explained by expansion of C 4 grasses at the expense of C 3 plants at low latitudes during glacial periods when atmospheric p CO 2 was lowered. C 4 dominance was overridingly influenced by global change in atmospheric p CO 2 despite the lowered summer rainfall.