Proposal for the Management of temporal and semantic components of Geographic Information

This poster raises the issue of a research work oriented to the storage, retrieval, representation and analysis of dynamic GI, taking into account The ultimate objective is the modelling and representation of the dynamic nature of geographic features, establishing mechanisms to store geometries enriched with a temporal structure (regardless of space) and a set of semantic descriptors detailing and clarifying the nature of the represented features and their temporality. the semantic, the temporal and the spatiotemporal components. We intend to define a set of methods, rules and restrictions for the adequate integration of these components into the primary elements of the GI: theme, location, time [1]. We intend to establish and incorporate three new structures (layers) into the core of data storage by using mark-up languages: a semantictemporal structure, a geosemantic structure, and an incremental spatiotemporal structure. Thus, data would be provided with the capability of pinpointing and expressing their own basic and temporal characteristics, enabling them to interact each other according to their context, and their time and meaning relationships that could be eventually established

Proposal for the Management of temporal and semantic components of Geographic Information

This poster raises the issue of a research work oriented to the storage, retrieval, representation and analysis of dynamic GI, taking into account the semantic, the temporal and the spatiotemporal components. We intend to define a set of methods, rules and restrictions for the adequate integration of these components into the primary elements of the GI: theme, location, time [1]. We intend to establish and incorporate three new structures (layers) into the core of data storage by using mark-up languages: a semantictemporal structure, a geosemantic structure, and an incremental spatiotemporal structure. The ultimate objective is the modelling and representation of the dynamic nature of geographic features, establishing mechanisms to store geometries enriched with a temporal structure (regardless of space) and a set of semantic descriptors detailing and clarifying the nature of the represented features and their temporality. Thus, data would be provided with the capability of pinpointing and expressing their own basic and temporal characteristics, enabling them to interact each other according to their context, and their time and meaning relationships that could be eventually established

Summer deficit irrigation strategies in hedgerow olive orchard cv. ‘Arbequina’: Effect on fruit characteristics and yield.

Maximum production in hedgerow olive orchards is likely not achieved with maximum evapotranspiration over the long-term. Thus, regulated deficit irrigation (RDI) should be considered as a management option. Four irrigation treatments were evaluated during the summer when olive is most drought resistant. Control (CON) was irrigated to maintain the rootzone close to field capacity. Severe water deficit was applied by irrigating 30% CON from end of fruit drop to end July (DI-J) and from end July until beginning of oil synthesis (DI-A). Less severe water deficit was applied during July and August (DI-JA) by irrigating 50% CON. Flowering, fruiting, abscission, fruit development, fresh and dry weight of fruits, and oil production were evaluated. There were not significant differences in number of buds initiated, number of fruits per inflorescence and fruit drop. Oil production was significantly different between irrigation treatments in all experimental years. CON produced more oil and fruit with higher oil% than DI-A and DI-JA. Oil production of DI-J was not significantly reduced compared to CON and oil% was greater. DI-J was the most effective RDI strategy; with 16% less applied water relative to CON average loss in oil production of 8% was not significantly different to CON. While DI-JA saved most water (27%), oil production was reduced by 15%. Greatest loss in oil production (21%) was observed in DI-A with water saving of 22%.

Integration of temporal and semantic components into the Geographic Information. Part II: Methodology

The overall objective of this research project is to enrich geographic data with temporal and semantic components in order to significantly improve spatio-temporal analysis of geographic phenomena. To achieve this goal, we intend to establish and incorporate three new layers (structures) into the core of the Geographic Information by using mark-up languages as well as defining a set of methods and tools for enriching the system to make it able to retrieve and exploit such layers (semantic-temporal, geosemantic, and incremental spatio-temporal). Besides these layers, we also propose a set of models (temporal and spatial) and two semantic engines that make the most of the enriched geographic data. The roots of the project and its definition have been previously presented in Siabato & Manso-Callejo 2011. In this new position paper, we extend such work by delineating clearly the methodology and the foundations on which we will base to define the main components of this research: the spatial model, the temporal model, the semantic layers, and the semantic engines. By putting together the former paper and this new work we try to present a comprehensive description of the whole process, from pinpointing the basic problem to describing and assessing the solution. In this new article we just mention the methods and the background to describe how we intend to define the components and integrate them into the GI.

Farm nitrogen balances in six European landscapes as an indicator for nitrogen losses and basis for improved management.

Improved management of nitrogen (N) in agriculture is necessary to achieve a sustainable balance between the production of food and other biomass, and the unwanted effects of N on water pollution, greenhouse gas emissions, biodiversity deterioration and human health. To analyse farm N-losses and the complex interactions within farming systems, efficient methods for identifying emissions hotspots and evaluating mitigation measures are therefore needed. The present paper aims to fill this gap at the farm and landscape scales. Six agricultural landscapes in Poland (PL), the Netherlands (NL), France (FR), Italy (IT), Scotland (UK) and Denmark (DK) were studied, and a common method was developed for undertaking farm inventories and the derivation of farm N balances, N surpluses and for evaluating uncertainty for the 222 farms and 11 440 ha of farmland included in the study. In all landscapes, a large variation in the farm N surplus was found, and thereby a large potential for reductions. The highest average N surpluses were found in the most livestock-intensive landscapes of IT, FR, and NL; on average 202 ± 28, 179 ± 63 and 178 ± 20 kg N ha−1 yr−1, respectively. All landscapes showed hotspots, especially from livestock farms, including a special UK case with large-scale landless poultry farming. Overall, the average N surplus from the land-based UK farms dominated by extensive sheep and cattle grazing was only 31 ± 10 kg N ha−1 yr−1, but was similar to the N surplus of PL and DK (122 ± 20 and 146 ± 55 kg N ha−1 yr−1, respectively) when landless poultry farming was included. We found farm N balances to be a useful indicator for N losses and the potential for improving N management. Significant correlations to N surplus were found, both with ammonia air concentrations and nitrate concentrations in soils and groundwater, measured during the period of N management data collection in the landscapes from 2007–2009. This indicates that farm N surpluses may be used as an independent dataset for validation of measured and modelled N emissions in agricultural landscapes. No significant correlation was found with N measured in surface waters, probably because of spatial and temporal variations in groundwater buffering and biogeochemical reactions affecting N flows from farm to surface waters. A case study of the development in N surplus from the landscape in DK from 1998–2008 showed a 22% reduction related to measures targeted at N emissions from livestock farms. Based on the large differences in N surplus between average N management farms and the most modern and N-efficient farms, it was concluded that additional N-surplus reductions of 25–50%, as compared to the present level, were realistic in all landscapes. The implemented N-surplus method was thus effective for comparing and synthesizing results on farm N emissions and the potentials of mitigation options. It is recommended for use in combination with other methods for the assessment of landscape N emissions and farm N efficiency, including more detailed N source and N sink hotspot mapping, measurements and modelling.

Effect of conservation tillage on soil physical properties, in-situ rainwater harvesting, and erosion control in arid and semi-arid regions

La escasez del agua en las regiones áridas y semiáridas se debe a la escasez de precipitaciones y la distribución desigual en toda la temporada, lo que hace de la agricultura de secano una empresa precaria. Un enfoque para mejorar y estabilizar el agua disponible para la producción de cultivos en estas regiones es el uso de tecnologías de captación de agua de lluvia in situ y su conservación. La adopción de los sistemas de conservación de la humedad del suelo in situ, tales como la labranza de conservación, es una de las estrategias para mejorar la gestión de la agricultura en zonas áridas y semiáridas. El objetivo general de esta tesis ha sido desarrollar una metodología de aplicación de labranza de depósito e investigar los efectos a corto plazo sobre las propiedades físicas del suelo de las diferentes prácticas de cultivo que incluyen labranza de depósito: (reservoir tillage, RT), la laboreo mínimo: (minimum tillage, MT), la no laboreo: (zero tillage, ZT) y laboreo convencional: (conventional tillage, CT) Así como, la retención de agua del suelo y el control de la erosión del suelo en las zonas áridas y semiáridas. Como una primera aproximación, se ha realizado una revisión profunda del estado de la técnica, después de la cual, se encontró que la labranza de depósito es un sistema eficaz de cosecha del agua de lluvia y conservación del suelo, pero que no ha sido evaluada científicamente tanto como otros sistemas de labranza. Los trabajos experimentales cubrieron tres condiciones diferentes: experimentos en laboratorio, experimentos de campo en una región árida, y experimentos de campo en una región semiárida. Para investigar y cuantificar el almacenamiento de agua a temperatura ambiente y la forma en que podría adaptarse para mejorar la infiltración del agua de lluvia recolectada y reducir la erosión del suelo, se ha desarrollado un simulador de lluvia a escala de laboratorio. Las características de las lluvias, entre ellas la intensidad de las precipitaciones, la uniformidad espacial y tamaño de la gota de lluvia, confirmaron que las condiciones naturales de precipitación son simuladas con suficiente precisión. El simulador fue controlado automáticamente mediante una válvula de solenoide y tres boquillas de presión que se usaron para rociar agua correspondiente a diferentes intensidades de lluvia. Con el fin de evaluar el método de RT bajo diferentes pendientes de superficie, se utilizaron diferentes dispositivos de pala de suelo para sacar un volumen idéntico para hacer depresiones. Estas depresiones se compararon con una superficie de suelo control sin depresión, y los resultados mostraron que la RT fue capaz de reducir la erosión del suelo y la escorrentía superficial y aumentar significativamente la infiltración. Luego, basándonos en estos resultados, y después de identificar la forma adecuada de las depresiones, se ha diseñado una herramienta combinada (sistema integrado de labranza de depósito (RT)) compuesto por un arado de una sola línea de chisel, una sola línea de grada en diente de pico, sembradora modificada, y rodillo de púas. El equipo fue construido y se utiliza para comparación con MT y CT en un ambiente árido en Egipto. El estudio se realizó para evaluar el impacto de diferentes prácticas de labranza y sus parámetros de funcionamiento a diferentes profundidades de labranza y con distintas velocidades de avance sobre las propiedades físicas del suelo, así como, la pérdida de suelo, régimen de humedad, la eficiencia de recolección de agua, y la productividad de trigo de invierno. Los resultados indicaron que la RT aumentó drásticamente la infiltración, produciendo una tasa que era 47.51% más alta que MT y 64.56% mayor que la CT. Además, los resultados mostraron que los valores más bajos de la escorrentía y pérdidas de suelos 4.91 mm y 0.65 t ha-1, respectivamente, se registraron en la RT, mientras que los valores más altos, 11.36 mm y 1.66 t ha-1, respectivamente, se produjeron en el marco del CT. Además, otros dos experimentos de campo se llevaron a cabo en ambiente semiárido en Madrid con la cebada y el maíz como los principales cultivos. También ha sido estudiado el potencial de la tecnología inalámbrica de sensores para monitorizar el potencial de agua del suelo. Para el experimento en el que se cultivaba la cebada en secano, se realizaron dos prácticas de labranza (RT y MT). Los resultados mostraron que el potencial del agua del suelo aumentó de forma constante y fue consistentemente mayor en MT. Además, con independencia de todo el período de observación, RT redujo el potencial hídrico del suelo en un 43.6, 5.7 y 82.3% respectivamente en comparación con el MT a profundidades de suelo (10, 20 y 30 cm, respectivamente). También se observaron diferencias claras en los componentes del rendimiento de los cultivos y de rendimiento entre los dos sistemas de labranza, el rendimiento de grano (hasta 14%) y la producción de biomasa (hasta 8.8%) se incrementaron en RT. En el experimento donde se cultivó el maíz en regadío, se realizaron cuatro prácticas de labranza (RT, MT, ZT y CT). Los resultados revelaron que ZT y RT tenían el potencial de agua y temperatura del suelo más bajas. En comparación con el tratamiento con CT, ZT y RT disminuyó el potencial hídrico del suelo en un 72 y 23%, respectivamente, a la profundidad del suelo de 40 cm, y provocó la disminución de la temperatura del suelo en 1.1 y un 0.8 0C respectivamente, en la profundidad del suelo de 5 cm y, por otro lado, el ZT tenía la densidad aparente del suelo y resistencia a la penetración más altas, la cual retrasó el crecimiento del maíz y disminuyó el rendimiento de grano que fue del 15.4% menor que el tratamiento con CT. RT aumenta el rendimiento de grano de maíz cerca de 12.8% en comparación con la ZT. Por otra parte, no hubo diferencias significativas entre (RT, MT y CT) sobre el rendimiento del maíz. En resumen, según los resultados de estos experimentos, se puede decir que mediante el uso de la labranza de depósito, consistente en realizar depresiones después de la siembra, las superficies internas de estas depresiones se consolidan de tal manera que el agua se mantiene para filtrarse en el suelo y por lo tanto dan tiempo para aportar humedad a la zona de enraizamiento de las plantas durante un período prolongado de tiempo. La labranza del depósito podría ser utilizada como un método alternativo en regiones áridas y semiáridas dado que retiene la humedad in situ, a través de estructuras que reducen la escorrentía y por lo tanto puede resultar en la mejora de rendimiento de los cultivos. ABSTRACT Water shortage in arid and semi-arid regions stems from low rainfall and uneven distribution throughout the season, which makes rainfed agriculture a precarious enterprise. One approach to enhance and stabilize the water available for crop production in these regions is to use in-situ rainwater harvesting and conservation technologies. Adoption of in-situ soil moisture conservation systems, such as conservation tillage, is one of the strategies for upgrading agriculture management in arid and semi-arid environments. The general aim of this thesis is to develop a methodology to apply reservoir tillage to investigate the short-term effects of different tillage practices including reservoir tillage (RT), minimum tillage (MT), zero tillage (ZT), and conventional tillage (CT) on soil physical properties, as well as, soil water retention, and soil erosion control in arid and semi-arid areas. As a first approach, a review of the state of the art has been done. We found that reservoir tillage is an effective system of harvesting rainwater and conserving soil, but it has not been scientifically evaluated like other tillage systems. Experimental works covered three different conditions: laboratory experiments, field experiments in an arid region, and field experiments in a semi-arid region. To investigate and quantify water storage from RT and how it could be adapted to improve infiltration of harvested rainwater and reduce soil erosion, a laboratory-scale rainfall simulator was developed. Rainfall characteristics, including rainfall intensity, spatial uniformity and raindrop size, confirm that natural rainfall conditions are simulated with sufficient accuracy. The simulator was auto-controlled by a solenoid valve and three pressure nozzles were used to spray water corresponding to different rainfall intensities. In order to assess the RT method under different surface slopes, different soil scooping devices with identical volume were used to create depressions. The performance of the soil with these depressions was compared to a control soil surface (with no depression). Results show that RT was able to reduce soil erosion and surface runoff and significantly increase infiltration. Then, based on these results and after selecting the proper shape of depressions, a combination implement integrated reservoir tillage system (integrated RT) comprised of a single-row chisel plow, single-row spike tooth harrow, modified seeder, and spiked roller was developed and used to compared to MT and CT in an arid environment in Egypt. The field experiments were conducted to evaluate the impact of different tillage practices and their operating parameters at different tillage depths and different forward speeds on the soil physical properties, as well as on runoff, soil losses, moisture regime, water harvesting efficiency, and winter wheat productivity. Results indicated that the integrated RT drastically increased infiltration, producing a rate that was 47.51% higher than MT and 64.56% higher than CT. In addition, results showed that the lowest values of runoff and soil losses, 4.91 mm and 0.65 t ha-1 respectively, were recorded under the integrated RT, while the highest values, 11.36 mm and 1.66 t ha -1 respectively, occurred under the CT. In addition, two field experiments were carried out in semi-arid environment in Madrid with barley and maize as the main crops. For the rainfed barley experiment, two tillage practices (RT, and MT) were performed. Results showed that soil water potential increased quite steadily and were consistently greater in MT and, irrespective of the entire observation period, RT decreased soil water potential by 43.6, 5.7, and 82.3% compared to MT at soil depths (10, 20, and 30 cm, respectively). In addition, clear differences in crop yield and yield components were observed between the two tillage systems, grain yield (up to 14%) and biomass yield (up to 8.8%) were increased by RT. For the irrigated maize experiment, four tillage practices (RT, MT, ZT, and CT) were performed. Results showed that ZT and RT had the lowest soil water potential and soil temperature. Compared to CT treatment, ZT and RT decreased soil water potential by 72 and 23% respectively, at soil depth of 40 cm, and decreased soil temperature by 1.1 and 0.8 0C respectively, at soil depth of 5 cm. Also, ZT had the highest soil bulk density and penetration resistance, which delayed the maize growth and decreased the grain yield that was 15.4% lower than CT treatment. RT increased maize grain yield about 12.8% compared to ZT. On the other hand, no significant differences among (RT, MT, and CT) on maize yield were found. In summary, according to the results from these experiments using reservoir tillage to make depressions after seeding, these depression’s internal surfaces are consolidated in such a way that the water is held to percolate into the soil and thus allowing time to offer moisture to the plant rooting zone over an extended period of time. Reservoir tillage could be used as an alternative method in arid and semi-arid regions and it retains moisture in-situ, through structures that reduce runoff and thus can result in improved crop yields.

Simulating potential growth and yield of oil palm (Elaeis guineensis) with PALMSIM: Model description, evaluation and application.

Reducing the gap between water-limited potential yield and actual yield in oil palm production systems through intensification is seen as an important option for sustainably increasing palm oil production. Simulation models can play an important role in quantifying water-limited potential yield, and therefore the scope for intensification, but no oil palm model exists that is both simple enough and at the same time incorporates sufficient plant physiological knowledge to be generally applicable across sites with different growing conditions. The objectives of this study therefore were to develop a model (PALMSIM) that simulates, on a monthly time step, the potential growth of oil palm as determined by solar radiation and to evaluate model performance against measured oil palm yields under optimal water and nutrient management for a range of sites across Indonesia and Malaysia. The maximum observed yield in the field matches the corresponding simulated yield for dry bunch weight with a RMSE of 1.7 Mg ha?1 year?1 against an observed yield of 18.8 Mg ha?1. Sensitivity analysis showed that PALMSIM is robust: simulated changes in yield caused by modifying the parameters by 10% are comparable to other tree crop model evaluations. While we acknowledge that, depending on the soils and climatic environment, yields may be often water limited, we suggest a relatively simple physiological approach to simulate potential yield, which can be usefully applied to high rainfall environments and is considered as a first step in developing an oil palm model that also simulates water-limited potential yield. To illustrate the application possibil- ities of the model, PALMSIM was used to create a potential yield map for Indonesia and Malaysia by sim- ulating the growth and yield at a resolution of 0.1?. This map of potential yield is considered as a first step towards a decision support tool that can identify potentially productive, but at the moment degraded sites in Indonesia and Malaysia. ?

A Genetic Analysis of Interactions with Spc110p Reveals Distinct Functions of Spc97p and Spc98p, Components of the Yeast γ-Tubulin Complex

The spindle pole body (SPB) in Saccharomyces cerevisiae functions as the microtubule-organizing center. Spc110p is an essential structural component of the SPB and spans between the central and inner plaques of this multilamellar organelle. The amino terminus of Spc110p faces the inner plaque, the substructure from which spindle microtubules radiate. We have undertaken a synthetic lethal screen to identify mutations that enhance the phenotype of the temperature-sensitive spc110–221 allele, which encodes mutations in the amino terminus. The screen identified mutations in SPC97 and SPC98, two genes encoding components of the Tub4p complex in yeast. The spc98–63 allele is synthetic lethal only with spc110 alleles that encode mutations in the N terminus of Spc110p. In contrast, the spc97 alleles are synthetic lethal with spc110 alleles that encode mutations in either the N terminus or the C terminus. Using the two-hybrid assay, we show that the interactions of Spc110p with Spc97p and Spc98p are not equivalent. The N terminus of Spc110p displays a robust interaction with Spc98p in two different two-hybrid assays, while the interaction between Spc97p and Spc110p is not detectable in one strain and gives a weak signal in the other. Extra copies of SPC98 enhance the interaction between Spc97p and Spc110p, while extra copies of SPC97 interfere with the interaction between Spc98p and Spc110p. By testing the interactions between mutant proteins, we show that the lethal phenotype in spc98–63 spc110–221 cells is caused by the failure of Spc98–63p to interact with Spc110–221p. In contrast, the lethal phenotype in spc97–62 spc110–221 cells can be attributed to a decreased interaction between Spc97–62p and Spc98p. Together, these studies provide evidence that Spc110p directly links the Tub4p complex to the SPB. Moreover, an interaction between Spc98p and the amino-terminal region of Spc110p is a critical component of the linkage, whereas the interaction between Spc97p and Spc110p is dependent on Spc98p.

Differential Expression and Functions of Cortical Myosin IIA and IIB Isotypes during Meiotic Maturation, Fertilization, and Mitosis in Mouse Oocytes and Embryos

To explore the role of nonmuscle myosin II isoforms during mouse gametogenesis, fertilization, and early development, localization and microinjection studies were performed using monospecific antibodies to myosin IIA and IIB isotypes. Each myosin II antibody recognizes a 205-kDa protein in oocytes, but not mature sperm. Myosin IIA and IIB demonstrate differential expression during meiotic maturation and following fertilization: only the IIA isoform detects metaphase spindles or accumulates in the mitotic cleavage furrow. In the unfertilized oocyte, both myosin isoforms are polarized in the cortex directly overlying the metaphase-arrested second meiotic spindle. Cortical polarization is altered after spindle disassembly with Colcemid: the scattered meiotic chromosomes initiate myosin IIA and microfilament assemble in the vicinity of each chromosome mass. During sperm incorporation, both myosin II isotypes concentrate in the second polar body cleavage furrow and the sperm incorporation cone. In functional experiments, the microinjection of myosin IIA antibody disrupts meiotic maturation to metaphase II arrest, probably through depletion of spindle-associated myosin IIA protein and antibody binding to chromosome surfaces. Conversely, the microinjection of myosin IIB antibody blocks microfilament-directed chromosome scattering in Colcemid-treated mature oocytes, suggesting a role in mediating chromosome–cortical actomyosin interactions. Neither myosin II antibody, alone or coinjected, blocks second polar body formation, in vitro fertilization, or cytokinesis. Finally, microinjection of a nonphosphorylatable 20-kDa regulatory myosin light chain specifically blocks sperm incorporation cone disassembly and impedes cell cycle progression, suggesting that interference with myosin II phosphorylation influences fertilization. Thus, conventional myosins break cortical symmetry in oocytes by participating in eccentric meiotic spindle positioning, sperm incorporation cone dynamics, and cytokinesis. Although murine sperm do not express myosin II, different myosin II isotypes may have distinct roles during early embryonic development.

Population based randomised study of uptake and yield of screening by flexible sigmoidoscopy compared with screening by faecal occult blood testing

Objectives: To compare the feasibility of mass screening by flexible sigmoidoscopy with screening by faecal occult blood testing (Haemoccult) and both tests combined.

Nitrogen management and the future of food: Lessons from the management of energy and carbon

The food system dominates anthropogenic disruption of the nitrogen cycle by generating excess fixed nitrogen. Excess fixed nitrogen, in various guises, augments the greenhouse effect, diminishes stratospheric ozone, promotes smog, contaminates drinking water, acidifies rain, eutrophies bays and estuaries, and stresses ecosystems. Yet, to date, regulatory efforts to limit these disruptions largely ignore the food system. There are many parallels between food and energy. Food is to nitrogen as energy is to carbon. Nitrogen fertilizer is analogous to fossil fuel. Organic agriculture and agricultural biotechnology play roles analogous to renewable energy and nuclear power in political discourse. Nutrition research resembles energy end-use analysis. Meat is the electricity of food. As the agriculture and food system evolves to contain its impacts on the nitrogen cycle, several lessons can be extracted from energy and carbon: (i) set the goal of ecosystem stabilization; (ii) search the entire production and consumption system (grain, livestock, food distribution, and diet) for opportunities to improve efficiency; (iii) implement cap-and-trade systems for fixed nitrogen; (iv) expand research at the intersection of agriculture and ecology, and (v) focus on the food choices of the prosperous. There are important nitrogen-carbon links. The global increase in fixed nitrogen may be fertilizing the Earth, transferring significant amounts of carbon from the atmosphere to the biosphere, and mitigating global warming. A modern biofuels industry someday may produce biofuels from crop residues or dedicated energy crops, reducing the rate of fossil fuel use, while losses of nitrogen and other nutrients are minimized.

Chaperonin-facilitated protein folding: optimization of rate and yield by an iterative annealing mechanism.

We develop a heuristic model for chaperonin-facilitated protein folding, the iterative annealing mechanism, based on theoretical descriptions of "rugged" conformational free energy landscapes for protein folding, and on experimental evidence that (i) folding proceeds by a nucleation mechanism whereby correct and incorrect nucleation lead to fast and slow folding kinetics, respectively, and (ii) chaperonins optimize the rate and yield of protein folding by an active ATP-dependent process. The chaperonins GroEL and GroES catalyze the folding of ribulose bisphosphate carboxylase at a rate proportional to the GroEL concentration. Kinetically trapped folding-incompetent conformers of ribulose bisphosphate carboxylase are converted to the native state in a reaction involving multiple rounds of quantized ATP hydrolysis by GroEL. We propose that chaperonins optimize protein folding by an iterative annealing mechanism; they repeatedly bind kinetically trapped conformers, randomly disrupt their structure, and release them in less folded states, allowing substrate proteins multiple opportunities to find pathways leading to the most thermodynamically stable state. By this mechanism, chaperonins greatly expand the range of environmental conditions in which folding to the native state is possible. We suggest that the development of this device for optimizing protein folding was an early and significant evolutionary event.