The Perseus Exobiology Mission on MIR Behaviour of Amino Acids and Peptides in Earth Orbit

Leucine, -methyl leucine and two peptides were exposed tospace conditions on board the MIR station during the Perseus-Exobiology mission. This long duration space mission was aimed at testing the delivery of prebiotic building blocks. Duringthis mission, two amino acids (leucine and -methyl leucine) and two peptides (leucine-diketopiperazine and trileucine thioethylester) were exposed in Earth orbit for threemonths. Basalt, clay and meteorite powder were also mixed with the samples in order to simulate the effects of potential meteorite protection. Analysis of the material after the flight did not reveal any racemization or polymerisation but did provideinformation regarding photochemical pathways for the degradationof leucine and of the tripeptide. Amino acids appeared to be moresensitive to UV radiation than peptides, the cyclic dipeptide being found to be as particularly resistant. Meteorite powder which exhibits the highest absorption in Vacuum UltraViolet (VUV)afforded the best protection to the organic molecules whereasmontmorillonite clay, almost transparent in VUV, was the leastefficient. By varying the thickness of the meteorite, we found that the threshold for efficient protection against radiation was about 5 m. The possible exogenous origin of biologicalbuilding blocks is discussed with respect to the stability to themolecules and the nature of the associated minerals.     

Survival of Microorganisms under the Extreme Conditions of the Atacama Desert

Spores of Bacillus subtilis, conidia of Aspergillus niger, versicolor and ochraceus andcells of Deinococcus radiodurans have been exposed in the dark at two locations (at about 23°S and 24°S) in the Atacama Desert for up to 15 months. B. subtilis spores (survival 15%) and A. niger conidia (survival 30%) outlived the other species. The survival of the conidiaand spores species was only slightly poorer than that of thecorresponding laboratory controls. However, the Deinococcus radiodurans cells did not survive the desertexposure, because they are readily inactivated at relativehumidities between 40 and 80% which typically occurduring desert nights. Cellular monolayers of the dry sporesand conidia have in addition been exposed to the full sunlight for up to several hours. The solar fluences causing 63% loss in viability (F₃₇-values) have been determined.These F₃₇-values are compared with those determined atother global locations such as Punta Arenas (53°S), Key Largo (25°N) or Mainz (50°N) during the same season. Thesolar UVB radiation kills even the most resistantmicroorganisms within a few hours due to DNA damages. Thedata are also discussed with respect to possible similaritiesbetween the climatic conditions of the recent Atacama Desertand the deserts of early Mars.     

Soil and water and its relationship to the origin of life

Soils of the terrestrial planets form at the boundaries between lithosphere, atmosphere and hydrosphere. Biogenesis occurred in these zones; thus, it is axiomatic that some, perhaps many, stages of biogenesis occurred in intimate association with the mineral constituents of soils. Because of a high surface to mass ratio and, consequently, a high surface reactivity, the layer lattice clay minerals are the most important of these. according to the geological record, clay minerals appeared very early on the primordial Earth. Recent investigations have confirmed their presence in carbonaceous meteorites and have indicated their occurrence on Mars. In this paper we collect pertinent physico-chemical data and summarize the organic reactions and interactions that are induced or catalyzed by clays. Many clay-organic reactions that do not occur readily at high water contents proceed rapidly at adsorbed water contents corresponding to surface coverages of one or two molecular layers. One or two monolayers of adsorbed water correspond to extremely dry on cold planetary environments. Some consequences of these factsvis à vis biogenesis on Mars are considered.     

Photobiology in space: An experiment on Spacelab I

The joint European/US Spacelab Mission I, scheduled for October 1983 for a 9 day lasting Earth-orbiting flight, provides a laboratory system for various disciplines of science, including exobiology. On the pallet, in the experiment ES 029 Microorganisms and Biomolecules in Space Hard Environment 316 dry samples ofBacillus subtilis spores will be exposed to space vacuum and/or selected wavelengths of solar UV radiation. After recovery action spectra of inactivation, mutation induction, reparability and photochemical damage in DNA and protein will be determined. The results will contribute to the understanding of the mechanism of the increased UV sensitivity of bacterial spores in vacuo and to a better assessment of the chance of survival of resistant life forms in space and of interplanetary transfer of life.     

Space Microbiology

Summary: The responses of microorganisms (viruses, bacterial cells, bacterial and fungal spores, and lichens) to selected factors of space (microgravity, galactic cosmic radiation, solar UV radiation, and space vacuum) were determined in space and laboratory simulation experiments. In general, microorganisms tend to thrive in the space flight environment in terms of enhanced growth parameters and a demonstrated ability to proliferate in the presence of normally inhibitory levels of antibiotics. The mechanisms responsible for the observed biological responses, however, are not yet fully understood. A hypothesized interaction of microgravity with radiation-induced DNA repair processes was experimentally refuted. The survival of microorganisms in outer space was investigated to tackle questions on the upper boundary of the biosphere and on the likelihood of interplanetary transport of microorganisms. It was found that extraterrestrial solar UV radiation was the most deleterious factor of space. Among all organisms tested, only lichens (Rhizocarpon geographicum and Xanthoria elegans) maintained full viability after 2 weeks in outer space, whereas all other test systems were inactivated by orders of magnitude. Using optical filters and spores of Bacillus subtilis as a biological UV dosimeter, it was found that the current ozone layer reduces the biological effectiveness of solar UV by 3 orders of magnitude. If shielded against solar UV, spores of B. subtilis were capable of surviving in space for up to 6 years, especially if embedded in clay or meteorite powder (artificial meteorites). The data support the likelihood of interplanetary transfer of microorganisms within meteorites, the so-called lithopanspermia hypothesis.     

Short-term effects of biological and physical forces on aggregate formation in soils with different clay mineralogy

The mechanisms resulting in the binding of primary soil particles into stable aggregates vary with soil parent material, climate, vegetation, and management practices. In this study, we investigated short-term effects of: (i) nutrient addition (Hoagland's solution), (ii) organic carbon (OC) input (wheat residue), (iii) drying and wetting action, and (iv) root growth, with or without dry–wet cycles, on aggregate formation and stabilization in three soils differing in weathering status and clay mineralogy. These soils included a young, slightly weathered temperate soil dominated by 2:1 (illite and chlorite) clay minerals; a moderately weathered soil with mixed [2:1 (vermiculite) and 1:1 (kaolinite)] clay mineralogy and oxides; and a highly weathered tropical soil dominated by 1:1 (kaolinite) clay minerals and oxides. Air-dried soil was dry sieved through a 250 m sieve to break up all macroaggregates and 100 g-subsamples were brought to field capacity and incubated for 42 days. After 14 and 42 days, aggregate stability was measured on field moist and air-dried soil, to determine unstable and stable aggregation respectively. In control treatments (i.e., without nutrient or organic matter addition, without roots and at constant moisture), the formation of unstable and stable macroaggregates (> 250 m) increased in the order: 2:1 clay soil < mixed clay soil < 1:1 clay soil. After 42 days of incubation, nutrient addition significantly increased both unstable and stable macroaggregates in the 2:1 and 1:1 clay soils. In all soils, additional OC input increased both unstable and stable macroaggregate formation. The increase in macroaggregation with OC input was highest for the mixed clay soil and lowest for the 1:1 clay soil. In general, drying and wetting cycles had a positive effect on the formation of macroaggregates. Root growth caused a decrease in unstable macroaggregates in all soils. Larger amounts of macroaggregates were found in the mixed clay and oxides soil when plants were grown under 50% compared to 100% field capacity conditions. We concluded that soils dominated by variable charge clay minerals (1:1 clays and oxides) have higher potential to form stable aggregates when OC concentrations are low. With additional OC inputs, the greatest response in stable macroaggregate formation occurred in soils with mixed mineralogy, which is probably a result of different binding mechanisms occurring: i.e., electrostatic bindings between 2:1 clays, 1:1 clays and oxides (i.e. mineral-mineral bindings), in addition to OM functioning as a binding agent between 2:1 and 1:1 clays.     

Responses ofBacillus subtilis spores to space environment: Results from experiments in space

Onboard of several spacecrafts (Apollo 16, Spacelab 1, LDEF), spores ofBacillus subtilis were exposed to selected parameters of space, such as space vacuum, different spectral ranges of solar UV-radiation and cosmic rays, applied separately or in combination, and we have studied their survival and genetic changes after retrieval. The spores survive extended periods of time in space — up to several years —, if protected against the high influx of solar UV-radiation. Water desorption caused by the space vacuum leads to structural changes of the DNA; the consequences are an increased mutation frequency and altered photobiological properties of the spores. UV-effects, such as killing and mutagenesis, are augmented, if the spores are in space vacuum during irradiation. Vacuum-specific photoproducts which are different from the spore photoproduct may cause the synergistic response of spores to the simultaneous action of UV and vacuum. The experiments provide an experimental test of certain steps of the panspermia hypothesis.Presented at the Session Water in the Solar System and Its Role in Exobiology during the 26th General Assembly of the European Geophysical Society, 22–26 April 1991 in Wiesbaden, Germany     

Mars Primordial Crust: Unique Sites for Investigating Proto-biologic Properties

The Martian meteorite collection suggests that intact outcrops or boulder-scale fragments of the 4.5 Ga Martian crust exist within tens of meters of the present day surface of Mars. Mars may be the only planet where such primordial crust samples, representing the first 100 Ma of a planet’s environment, are available. The primordial crust has been destroyed on Earth by plate tectonics and other geological phenomena and is buried on the Moon under hundreds or thousands of meters of megaregoltih. Early Mars appears to have been remarkably similar to early Earth, and samples of rock from the first few Ma or first 100 Ma may reveal “missing link” proto-biological forms that could shed light on the transition from abiotic organic chemistry to living cells. Such organic snapshots of nascent life are unlikely to be found on Earth. Presented at: National Workshop on Astrobiology: Search for Life in the Solar System, Capri, Italy, 26 to 28 October, 2005.     

Photobiology in space: an experiment on Spacelab I

The joint European/US Spacelab Mission I, scheduled for October 1983 for a 9 day lasting Earth-orbiting flight, provides a laboratory system for various disciplines of science, including exobiology. On the pallet, in the experiment ES 029 "Microorganisms and Biomolecules in Space Hard Environment" 316 dry samples of Bacillus subtilis spores will be exposed to space vacuum and/or selected wavelenghs of solar UV radiation. After recovery action spectra of inactivation, mutation induction, reparability and photochemical damage in DNA and protein will be determined. The results will contribute to the understanding of the mechanism of the increased UV sensitivity of bacterial spores in vacuo and to a better assessment of the chance of survival of resistant life forms in space and of interplanetary transfer of life.     

Radiation-dependent limit for the viability of bacterial spores in halite fluid inclusions and on Mars

When claims for the long-term survival of viable organisms are made, either within terrestrial minerals or on Mars, considerations should be made of the limitations imposed by the naturally occurring radiation dose to which they have been exposed. We investigated the effect of ionizing radiation on different bacterial spores by measuring the inactivation constants for B. subtilis and S. marismortui spores in solution as well as for dry spores of B. subtilis and B. thuringiensis. S. marismortui is a halophilic spore that is genetically similar to the recently discovered 2-9-3 bacterium from a halite fluid inclusion, claimed to be 250 million years old (Vreeland et al., Nature 407, 897-900, 2000). B. thuringiensis is a soil bacterium that is genetically similar to the human pathogens B. anthracis and B. cereus (Helgason et al., Appl. Environ. Microbiol. 66, 2627-2630, 2000). To relate the inactivation constant to some realistic environments, we calculated the radiation regimen in a halite fluid inclusion and in the Martian subsurface over time. Our conclusion is that the ionizing dose of radiation in those environments limits the survival of viable bacterial spores over long periods. In the absence of an active repair mechanism in the dormant state, the long-term survival of spores is limited to less than 109 million years in halite fluid inclusions, to 100 to 160 million years in the Martian subsurface below 3 m, and to less than 600,000 years in the uppermost meter of Mars.     

The solar UV environment and bacterial spore UV resistance: considerations for Earth-to-Mars transport by natural processes and human spaceflight

The environment in space and on planets such as Mars can be lethal to microorganisms because of the high vacuum and high solar radiation flux, in particular UV radiation, in such environments. Spores of various Bacillus species are among the organisms most resistant to the lethal effects of high vacuum and UV radiation, and as a consequence are of major concern for planetary contamination via unmanned spacecraft or even natural processes. This review focuses on the spores of various Bacillus species: (i) their mechanisms of UV resistance; (ii) their survival in unmanned spacecraft, space flight and simulated space flight and Martian conditions; (iii) the UV flux in space and on Mars; (iv) factors affecting spore survival in such high UV flux environments.     

The survival of micromycetes exposed to space conditions

Original data on the survival of fungal spores exposed to space conditions are presented. The experiment was carried out on the Earth-orbiting Russian satellite Foton-M4. The flight duration of the satellite was 45 days. Thirteen fungal species (hyaline as well as pigmented) from 10 genera recovered from destructed stone materials were studied. Sterile quartz sand was inoculated by the fungal spores and was placed into Eppendorf tubes. During the space flight, the Eppendorf tubes with fungal spores were kept inside the Foton descent capsule in the “Biokont” containers and on the external surface of the capsule in the “Exobiofrost” containers exposed to the open space as well. Spores of ten species (77% of all tested species), i.e. Acremonium charticola, Aspergillus niger, Aspergillus versicolor, Chaetomium globosum, Cladosporium sphaerospermum, Penicillium chrysogenum, Penicillium verrucosum, Purpureocillium lilacinum, Sarocladium kiliense, and Trichoderma harzianum, survived after the flight both inside and outside the descent capsule. Only three species (23% of all tested species), i.e. Acremonium furcatum, Engyodontium album and Verticillium zaregamsianum, failed to survive outside as well as inside the capsule. Spore viability differed depending on the fungal species. Thus, spores of some fungal species are able to survive under the complex of stress factors such as low temperature values, radiation, etc. We have shown that micromycetes can be used as a model group for study of eukaryotic organisms’ resistance to stress factors, due to their high tolerance not only to extreme terrestrial environments, but to the extraterrestrial ones as well.     

Soil and water and its relationship to the origin of life.

Soils of the terrestrial planets form at the boundaries between lithosphere, atmosphere and hydrosphere. Biogenesis occurred in these zones; thus, it is axiomatic that some, perhaps many, stages of biogensis occurred in intimate association with the mineral constituents of soils. Because of a high surface to mass ration and, consequently, a high surface reactivity, the layer lattice clay minerals are the most important of these. According to the geological record, clay minerals appeared very early on the primordial Earth. Recent investigations have confirmed their presence in carbonaceous meteorites and have indicated their occurrence on Mars. In this paper we collect pertinent physico-chemical data and summarize the organic reactions and interactions that are induced or catalyzed by clays. Many clay-organic reactions that do not occur readily at high water contents proceed rapidly at adsorbed water contents corresponding to surface coverages of one or two molecular layers. One or two monolayers of adsorbed water correspond to extremely dry or cold planetary environments. Some consequences of these facts vis á vis biogenesis on Mars are considered.     

The phytopathogen powdery mildew affects food-searching behavior and survival of <Emphasis Type="Italic">Coccinella septempunctata</Emphasis>

The diet of entomophagous coccinellids is mainly based on aphids and other food sources such as pollen, nectar, or fungal spores. Knowledge of their foraging behavior on plants infected by powdery mildew and their survival on fungal spores is currently limited. In this study, we investigated the olfactory response of Coccinella septempunctata to odor emission of barley plants infected by powdery mildew and their survival on fungal spores in the presence or absence of aphids. Odors released by powdery-mildew infected plants were more attractive for ladybirds compared to those of uninfected controls. After 3 days, the survival rate of ladybirds feeding only on powdery-mildew spores was less than 50%, while for ladybirds feeding exclusively on Rhopalosiphum padi aphids, the survival rate was close to 90%. After 15 days, the highest survival rate (almost 80%) was observed for ladybirds feeding on plants with both aphids and powdery mildew. Molecular analyses confirmed the presence of fungal spores in ladybird guts when feeding either on powdery mildew or on a mixed diet. Our results provide new insights into foraging behavior of entomophagous coccinellids revealing the potential of powdery mildew to be utilized as important non-essential food in a mixed diet, but also its lethal effect if consumed alone.     

Soil physical behaviour and crop responses to tillage in lowland rice soils of varying clay content

The influence of various tillage methods on two wetland rice soils in the Philippines is reported. The soils differed principally in clay content, 38% for the clay loam (clayey, mixed isohyperthermic Entic Hapludoll) while 56% for the clay (clayey, mixed noncalcareous, isohyperthermic Andaqueptic Haplaquoll). This had a marked effect on their response to tillage and varying water regime. The clay soil, under field conditions, showed little change in pore size distribution or soil water behaviour with different tillage methods. Crop (Rice, Oryza sativa L., var. IR20) yields were unaffected by tillage.In contrast, tillage effects were very marked in the clay loam soil, which consisted of a greenhouse and a field trial. In the greenhouse, which experienced severe dry periods, wet tillage not only increased the moisture retentivity but also the soil impedance at soil matric potential ()<–0.01 MPa. Seasonal average was <–1 MPa. Root length density decreased by 39% with dry tillage and by 56% with wet tillage compared with zero tillage. Grain yield however, did not vary with soil treatment. In the field, which experienced moderate dry spells, varied between –0.13 and –0.48 MPa. Root length density was significantly reduced at soil impedance >0.75 MPa. Wet tillage increased soil moisture storage which minimized the soil impedance during the dry cycle more effectively than did dry tillage. The crop performed best under wet tillage and least under zero tillage. Wet tillage in this soil was more effective under moderate than under severe water stress conditions.     

Glomeromycotan mycorrhizal fungi from tropical Australia III. Measuring diversity in natural and disturbed habitats

Aims and Background

The aim was to investigate the diversity and distribution of Glomeromycotan fungi forming arbuscular mycorrhizal associations (AMF) in undisturbed and disturbed habitats in the vicinity of Kakadu National Park in tropical Australia. This is a tropical region with a 7–9 month dry season and a monsoonal wet season. Complimentary methods of fungus detection were used to investigate the diversity and relative dominance of AMF at a regional scale.

Methods

Soils were sampled from 32 sites, representing eucalypt savanna woodlands, wetlands, sandstone escarpment, rainforest, and disturbed mine waste rock dumps (overburden or spoil). Populations of AMF were identified and quantified using spores from soil. Morphology patterns of fungi colonising bait plant roots were examined and isolates were obtained by four complimentary pot-culturing methods.

Results

Different methods of detecting fungi produced different answers about which AMF were most important in the tested soils. In particular, spore surveys apparently underestimated the importance of Glomus species and overestimated the activity of Acaulospora species with numerous small spores, while calculated spore biovolumes overestimated the importance of Scutellospora and Gigaspora species with large spores, relative to inoculum levels of these fungus categories measured in bioassays. Spore surveys revealed 15 species of fungi and 8 additional fungi were recovered from the same soil samples using pot-culture isolation methods. Pot-cultures were especially important for detecting Glomus species that had high inoculum levels, but rarely produced spores in soils. Spores of AMF increased in abundance as vegetation developed in mine habitats reaching a peak that was higher than in undisturbed plant communities. Spore numbers (but not biovolumes) were well correlated with bioassay measurements of inoculum levels.

Conclusions

Most AMF species were widespread, but several were restricted to disturbed habitats or wetland soils. Undisturbed sites had a substantially higher diversity of AMF than partially vegetated mine waste rock dumps. It is recommended that AMF population surveys should not be based entirely on spore occurrence data, to avoid overlooking important fungi that sporulate infrequently. These fungi could be detected by bioassays or pot culture isolation from soil. Major variations in the detectability of AMF correspond to different life history strategies and can mask variations in their abundance.     

LIFE Experiment: Isolation of Cryptoendolithic Organisms from Antarctic Colonized Sandstone Exposed to Space and Simulated Mars Conditions on the International Space Station

Desiccated Antarctic rocks colonized by cryptoendolithic communities were exposed on the International Space Station (ISS) to space and simulated Mars conditions (LiFE-Lichens and Fungi Experiment). After 1.5?years in space samples were retrieved, rehydrated and spread on different culture media. Colonies of a green alga and a pink-coloured fungus developed on Malt-Agar medium; they were isolated from a sample exposed to simulated Mars conditions beneath a 0.1?%?T Suprasil neutral density filter and from a sample exposed to space vacuum without solar radiation exposure, respectively. None of the other flight samples showed any growth after incubation. The two organisms able to grow were identified at genus level by Small SubUnit (SSU) and Internal Transcribed Spacer (ITS) rDNA sequencing as Stichococcus sp. (green alga) and Acarospora sp. (lichenized fungal genus) respectively. The data in the present study provide experimental information on the possibility of eukaryotic life transfer from one planet to another by means of rocks and of survival in Mars environment.     

Modeling soil–root water transport and competition for single and mixed crops

A knowledge of above and below ground plant interactions for water is essential to understand the performance of intercropped systems. In this work, root water potential dynamics and water uptake partitioning were compared between single crops and intercrops, using a simulation model. Four root maps having 498, 364, 431 and 431 soil-root contacts were used. In the first and second cases, single crops with deep and surface roots were considered, whereas in the third and fourth cases, roots of two mixed crops were simultaneously considered with different row spacing (40 cm and 60 cm). Two soils corresponding to a clay and a silty clay loam were used in the calculations. A total maximum evapotranspiration of 6 mm d^-1 for both single or mixed crops was considered, for the mixed crops however, two transpiration distributions between the crops were analyzed (3:3 mm d^-1, or 4:2 mm d^-1 for each crop, respectively). The model was based on a previous theoretical framework applied to single or intercropped plants having spatially distributed roots in a two-dimensional domain. Although water stress occurred more rapidly in the loam than in the clay, due to the rapid decrease of the soil water reserve in the loam, the role of the root arrangement appeared to be crucial for water availability. Interactions between the distribution of transpiration among mixed crops and the architecture of the root systems which were in competition led to water movements from zones with one plant to another, or vice versa, which corresponded to specific competition or facilitation effects. Decreasing the distances between roots may increase competition for water, although it may determine greater water potential gradients in the soil that increase lateral or vertical water fluxes in the soil profile. The effects of the root competition on water uptake were quite complicated, depending on both environmental conditions, soil hydrodynamic properties, and time scales. Although some biological adaptive mechanisms were disregarded in the analysis, the physically 2-D based model may be considered as a tool to study the exploitation of environmental heterogeneity at microsite scales.     

<Emphasis Type="Italic">Rhizophagus irregularis</Emphasis> MUCL 41833 can colonize and improve P uptake of <Emphasis Type="Italic">Plantago lanceolata</Emphasis> after exposure to ionizing gamma radiation in root organ culture

Long-lived radionuclides such as ⁹⁰Sr and ¹³⁷Cs can be naturally or accidentally deposited in the upper soil layers where they emit β/γ radiation. Previous studies have shown that arbuscular mycorrhizal fungi (AMF) can accumulate and transfer radionuclides from soil to plant, but there have been no studies on the direct impact of ionizing radiation on AMF. In this study, root organ cultures of the AMF Rhizophagus irregularis MUCL 41833 were exposed to 15.37, 30.35, and 113.03 Gy gamma radiation from a ¹³⁷Cs source. Exposed spores were subsequently inoculated to Plantago lanceolata seedlings in pots, and root colonization and P uptake evaluated. P. lanceolata seedlings inoculated with non-irradiated AMF spores or with spores irradiated with up to 30.35 Gy gamma radiation had similar levels of root colonization. Spores irradiated with 113.03 Gy gamma radiation failed to colonize P. lanceolata roots. P content of plants inoculated with non-irradiated spores or of plants inoculated with spores irradiated with up to 30.35 Gy gamma radiation was higher than in non-mycorrhizal plants or plants inoculated with spores irradiated with 113.03 Gy gamma radiation. These results demonstrate that spores of R. irregularis MUCL 41833 are tolerant to chronic ionizing radiation at high doses.     

Growth and phosphorus nutrition of rice when inorganic fertiliser application is partly replaced by straw under varying moisture availability in sandy and clay soils

Background and aims

The combined effects of (1) reduced soil moisture availability, (2) reduced application of inorganic fertilisers while incorporating straw, (3) soil type, and their effects on growth, root system plasticity, phosphorus (P) nutrition of rice, and soil P dynamics are poorly known, but very important when aiming to increase the efficiency of water and P use.

Methods

Using large pots a three-factor factorial experiment was conducted with two moisture treatments (i.e. continuous flooding, and draining of top soil after flowering while subsoil was kept moist through capillary action), three fertilisation treatments; with (P1) and without (P0) applications of inorganic P fertilisers, and 25 % of inorganic fertilisers reduced while incorporating straw (5 t ha^?1), and soil type (i.e. clay and sandy soils with 15 and 9 mg P kg^?1 soil, respectively in P0). Shoot and root growth, root system plasticity, P nutrient status and soil P dynamics were measured.

Key results

Straw incorporation with reduced inorganic fertiliser application ensured a higher shoot dry weight and yield only in flooded clay soil as compared with P0 and P1, and a similar shoot dry weight and yield to P1 under drained clay soil. A positive growth response was facilitated by an increased water-use efficiency and rate of photosynthesis in shoots, and increased root system plasticity through the production of greater root length, more roots in deep soil layers, and an increased fraction of fine roots. Straw enhanced P extractability in soil. Drained soil reduced P uptake (15–45 %) and increased P-use efficiency. In addition to the re-translocation of P from senescing leaves and stems under both moisture conditions, the P concentration in green leaves under drained condition was also reduced (41–72 %).

Conclusion

Growth benefits of straw incorporation were observed in clay soil under both moisture conditions, and this was facilitated by the improved P availability, increased P uptake, and greater root system plasticity with the production of deeper and finer roots, compared with that in sandy soil, and inorganic fertiliser applications alone. As P uptake was reduced under drained soil, P re-translocation and % P allocated to panicles increased.