Geochemistry of Ontong Java Plateau basalts

Basement rocks from the Ontong Java Plateau are tholeiitic basalts that appear to record very high degrees of partial melting, much like those found today in the vicinity of Iceland. They display a limited range of incompatible element and isotopic variation, but small differences are apparent between sampled sites and between upper and lower groups of flows at Ocean Drilling Program Site 807.40Ar-39Ar ages of lavas from Site 807 and Deep Sea Drilling Project Site 289 are indistinguishable about an early Aptian mean of 122 Ma (as are preliminary data for the island of Malaita at the southern edge of the plateau), indicating that plateau-building eruptions ended more or less simultaneously at widely separated locations. Pb-Nd-Sr isotopes for lavas from Sites 289, 803, and 807, as well as southern Malaita, reflect a hotspot-like source with epsilon-Nd(T) = +4.0 to +6.3, (87Sr/86Sr)T = 0.70423-0.70339, and 206Pb/204Pb = 18.245-18.709 and possessing consistently greater 208Pb/204Pb for a given 206Pb/204Pb than Pacific MORB. The combination of hotspot-like mantle source, very high degrees of melting, and lack of a discernible age progression is best explained if the bulk of the plateau was constructed rapidly above a surfacing plume head, possibly that of the Louisville hotspot. Basalt and feldspar separates indicate a substantially younger age of ~90 Ma for basement at Site 803; in addition, volcaniclastic layers of mid-Cenomanian through Coniacian age occur at DSDP Site 288, and beds of late Aptian-Albian age are found at Site 289. Therefore, at least some volcanism continued on the plateau for 30 m.y. or more. The basalts at Site 803 are chemically and isotopically very similar to those at the ~122 Ma sites, suggesting that hot plume-type mantle was present beneath the plateau for an extended period or at two different times. Surviving seamounts of the Louisville Ridge formed between 70 and 0 Ma have much higher 206Pb/204Pb than any of the plateau basalts. Thus, assuming the Louisville hotspot was the source of the plateau lavas, a change in the hotspot's isotopic composition may have occurred between roughly 70 and 90 Ma; such a change may have accompanied the plume-head to plume-tail transition. Similar shifts from early, lower 206Pb/204Pb to subsequently higher 206Pb/204Pb values are found in several other oceanic plateau-hotspot and continental flood basalt-hotspot systems, and could reflect either a reduction in the supply of low 206Pb/204Pb mantle or an inability of some off-ridge plume-tails to melt refractory low 206Pb/204Pb material.

Modelización de series temporales complejas: de las redes de telecomunicación a los mercados financieros

Este trabajo aborda el problema de modelizar sistemas din´amicos reales a partir del estudio de sus series temporales, usando una formulaci´on est´andar que pretende ser una abstracci´on universal de los sistemas din´amicos, independientemente de su naturaleza determinista, estoc´astica o h´ıbrida. Se parte de modelizaciones separadas de sistemas deterministas por un lado y estoc´asticos por otro, para converger finalmente en un modelo h´ıbrido que permite estudiar sistemas gen´ericos mixtos, esto es, que presentan una combinaci´on de comportamiento determinista y aleatorio. Este modelo consta de dos componentes, uno determinista consistente en una ecuaci´on en diferencias, obtenida a partir de un estudio de autocorrelaci´on, y otro estoc´astico que modeliza el error cometido por el primero. El componente estoc´astico es un generador universal de distribuciones de probabilidad, basado en un proceso compuesto de variables aleatorias, uniformemente distribuidas en un intervalo variable en el tiempo. Este generador universal es deducido en la tesis a partir de una nueva teor´ıa sobre la oferta y la demanda de un recurso gen´erico. El modelo resultante puede formularse conceptualmente como una entidad con tres elementos fundamentales: un motor generador de din´amica determinista, una fuente interna de ruido generadora de incertidumbre y una exposici´on al entorno que representa las interacciones del sistema real con el mundo exterior. En las aplicaciones estos tres elementos se ajustan en base al hist´orico de las series temporales del sistema din´amico. Una vez ajustados sus componentes, el modelo se comporta de una forma adaptativa tomando como inputs los nuevos valores de las series temporales del sistema y calculando predicciones sobre su comportamiento futuro. Cada predicci´on se presenta como un intervalo dentro del cual cualquier valor es equipro- bable, teniendo probabilidad nula cualquier valor externo al intervalo. De esta forma el modelo computa el comportamiento futuro y su nivel de incertidumbre en base al estado actual del sistema. Se ha aplicado el modelo en esta tesis a sistemas muy diferentes mostrando ser muy flexible para afrontar el estudio de campos de naturaleza dispar. El intercambio de tr´afico telef´onico entre operadores de telefon´ıa, la evoluci´on de mercados financieros y el flujo de informaci´on entre servidores de Internet son estudiados en profundidad en la tesis. Todos estos sistemas son modelizados de forma exitosa con un mismo lenguaje, a pesar de tratarse de sistemas f´ısicos totalmente distintos. El estudio de las redes de telefon´ıa muestra que los patrones de tr´afico telef´onico presentan una fuerte pseudo-periodicidad semanal contaminada con una gran cantidad de ruido, sobre todo en el caso de llamadas internacionales. El estudio de los mercados financieros muestra por su parte que la naturaleza fundamental de ´estos es aleatoria con un rango de comportamiento relativamente acotado. Una parte de la tesis se dedica a explicar algunas de las manifestaciones emp´ıricas m´as importantes en los mercados financieros como son los “fat tails”, “power laws” y “volatility clustering”. Por ´ultimo se demuestra que la comunicaci´on entre servidores de Internet tiene, al igual que los mercados financieros, una componente subyacente totalmente estoc´astica pero de comportamiento bastante “d´ocil”, siendo esta docilidad m´as acusada a medida que aumenta la distancia entre servidores. Dos aspectos son destacables en el modelo, su adaptabilidad y su universalidad. El primero es debido a que, una vez ajustados los par´ametros generales, el modelo se “alimenta” de los valores observables del sistema y es capaz de calcular con ellos comportamientos futuros. A pesar de tener unos par´ametros fijos, la variabilidad en los observables que sirven de input al modelo llevan a una gran riqueza de ouputs posibles. El segundo aspecto se debe a la formulaci´on gen´erica del modelo h´ıbrido y a que sus par´ametros se ajustan en base a manifestaciones externas del sistema en estudio, y no en base a sus caracter´ısticas f´ısicas. Estos factores hacen que el modelo pueda utilizarse en gran variedad de campos. Por ´ultimo, la tesis propone en su parte final otros campos donde se han obtenido ´exitos preliminares muy prometedores como son la modelizaci´on del riesgo financiero, los algoritmos de routing en redes de telecomunicaci´on y el cambio clim´atico. Abstract This work faces the problem of modeling dynamical systems based on the study of its time series, by using a standard language that aims to be an universal abstraction of dynamical systems, irrespective of their deterministic, stochastic or hybrid nature. Deterministic and stochastic models are developed separately to be merged subsequently into a hybrid model, which allows the study of generic systems, that is to say, those having both deterministic and random behavior. This model is a combination of two different components. One of them is deterministic and consisting in an equation in differences derived from an auto-correlation study and the other is stochastic and models the errors made by the deterministic one. The stochastic component is an universal generator of probability distributions based on a process consisting in random variables distributed uniformly within an interval varying in time. This universal generator is derived in the thesis from a new theory of offer and demand for a generic resource. The resulting model can be visualized as an entity with three fundamental elements: an engine generating deterministic dynamics, an internal source of noise generating uncertainty and an exposure to the environment which depicts the interactions between the real system and the external world. In the applications these three elements are adjusted to the history of the time series from the dynamical system. Once its components have been adjusted, the model behaves in an adaptive way by using the new time series values from the system as inputs and calculating predictions about its future behavior. Every prediction is provided as an interval, where any inner value is equally probable while all outer ones have null probability. So, the model computes the future behavior and its level of uncertainty based on the current state of the system. The model is applied to quite different systems in this thesis, showing to be very flexible when facing the study of fields with diverse nature. The exchange of traffic between telephony operators, the evolution of financial markets and the flow of information between servers on the Internet are deeply studied in this thesis. All these systems are successfully modeled by using the same “language”, in spite the fact that they are systems physically radically different. The study of telephony networks shows that the traffic patterns are strongly weekly pseudo-periodic but mixed with a great amount of noise, specially in the case of international calls. It is proved that the underlying nature of financial markets is random with a moderate range of variability. A part of this thesis is devoted to explain some of the most important empirical observations in financial markets, such as “fat tails”, “power laws” and “volatility clustering”. Finally it is proved that the communication between two servers on the Internet has, as in the case of financial markets, an underlaying random dynamics but with a narrow range of variability, being this lack of variability more marked as the distance between servers is increased. Two aspects of the model stand out as being the most important: its adaptability and its universality. The first one is due to the fact that once the general parameters have been adjusted , the model is “fed” on the observable manifestations of the system in order to calculate its future behavior. Despite the fact that the model has fixed parameters the variability in the observable manifestations of the system, which are used as inputs of the model, lead to a great variability in the possible outputs. The second aspect is due to the general “language” used in the formulation of the hybrid model and to the fact that its parameters are adjusted based on external manifestations of the system under study instead of its physical characteristics. These factors made the model suitable to be used in great variety of fields. Lastly, this thesis proposes other fields in which preliminary and promising results have been obtained, such as the modeling of financial risk, the development of routing algorithms for telecommunication networks and the assessment of climate change.

Electrodynamic Tethers For Planetary And De-orbiting Missions

Nuevas aplicaciones tecnológicas y científicas mediante amarras electrodinámicas son analizadas para misiones planetarias. i) Primero, se considera un conjunto de amarras cilíndricas en paralelo (veleros electrosolares) para una misión interplanetaria. Los iones provenientes del viento solar son repelidos por el alto potencial de dichas amarras generando empuje sobre el velero. Para conocer el intercambio de momento que provocan los iones sobre las amarras se ha considerado un modelo de potencial estacionario. Se ha analizado la transferencia orbital de la Tierra a Júpiter siguiendo un método de optimización de trayectoria indirecto. ii) Una vez que el velero se encuentra cerca de Júpiter, se ha considerado el despliegue de una amarra para diferentes objetivos científicos. iia) Una amarra podría ser utilizada para diagnóstico de plasmas, al ser una fuente efectiva de ondas, y también como un generador de auroras artificiales. Una amarra conductora que orbite en la magnetosfera jovial es capaz de producir ondas. Se han analizado las diferentes ondas radiadas por un conductor por el que circula una corriente constante que sigue una órbita polar de alta excentricidad y bajo apoápside, como ocurre en la misión Juno de la NASA. iib) Además, se ha estudiado una misión tentativa que sigue una órbita ecuatorial (LJO) por debajo de los intensos cinturones de radiación. Ambas misiones requiren potencia eléctrica para los sistemas de comunicación e instrumentos científicos. Las amarras pueden generar potencia de manera más eficiente que otros sistemas que utlizan paneles solares o sistemas de potencia de radioisótopos (RPS). La impedancia de radiación es necesaria para determinar la corriente que circula por todo el circuito de la amarra. En un modelo de plasma frío, la radiación ocurre principalmente en los modos de Alfven y magnetosónica rápida, mostrando un elevado índice de refracción. Se ha estudiado la impedancia de radiación en amarras con recubrimiento aislante para los dos modos de radiación y cada una de las misiones. A diferencia del caso ionosférico terrestre, la baja densidad y el intenso campo magnético que aparecen en el entorno de Júpiter consiguen que la girofrecuencia de los electrones sea mucho mayor que la frecuencia del plasma; esto hace que el espectro de potencia para cada modo se modifique substancialmente, aumentando la velocidad de Alfven. Se ha estimado también la impedancia de radiación para amarras sin aislante conductor. En la misión LJO, un vehículo espacial bajando lentamente la altitud de su órbita permitiría estudiar la estructura del campo magnético y composición atmosférica para entender la formación, evolución, y estructura de Júpiter. Adicionalmente, si el contactor (cátodo) se apaga, se dice que la amarra flota eléctricamente, permitiendo emisión de haz de electrones que generan auroras. El continuo apagado y encendido produce pulsos de corriente dando lugar a emisiones de señales, que pueden ser utilizadas para diagnóstico del plasma jovial. En Órbita Baja Jovial, los iones que impactan contra una amarra polarizada negativamente producen electrones secundarios, que, viajando helicoidalmente sobre las líneas de campo magnético de Júpiter, son capaces de alcanzar su atmósfera más alta, y, de esta manera, generar auroras. Se han identificado cuáles son las regiones donde la amarra sería más eficiente para producir auroras. iic) Otra aplicación científica sugerida para la misión LJO es la detección de granos cargados que orbitan cerca de Júpiter. Los electrones de alta energía en este ambiente pueden ser modelados por una distribucción no Maxwelliana conocida como distribución kappa. En escenarios con plasmas complejos, donde los campos eléctricos en Júpiter pueden acelerar las cargas hasta velocidades que superen la velocidad térmica, este tipo de distribuciones son muy útiles. En este caso las colas de las distribuciones de electrones siguen una ley de potencias. Se han estudiado las fluctuaciones de granos cargados para funciones de distribución kappa. iii) La tesis concluye con el análisis para deorbitar satélites con amarras electrodinámicas que siguen una Órbita Baja Terrestre (LEO). Una amarra debe presentar una baja probabilidad de corte por pequeño debris y además debe ser suficientemente ligero para que el cociente entre la masa de la amarra y el satélite sea muy pequeño. En este trabajo se estiman las medidas de la longitud, anchura y espesor que debe tener una amarra para minimizar el producto de la probabilidad de corte por el cociente entre las masas de la amarra y el satélite. Se presentan resultados preliminares del diseño de una amarra con forma de cinta para deorbitar satélites relativamente ligeros como Cryosat y pesados como Envisat. Las misiones espaciales a planetas exteriores y en el ámbito terrestre plantean importantes retos científico-tecnológicos que deben ser abordados y solucionados. Por ello, desde el inicio de la era espacial se han diseñando novedosos métodos propulsivos, sistemas de guiado, navegación y control más robustos, y nuevos materiales para mejorar el rendimiento de los vehículos espaciales (SC). En un gran número de misiones interplanetarias y en todas las misiones a planetas exteriores se han empleado sistemas de radioisótopos (RPS) para generar potencia eléctrica en los vehículos espaciales y en los rovers de exploración. Estos sistemas emplean como fuente de energía el escaso y costoso plutonio-238. La NASA, por medio de un informe de la National Academy of Science (5 de Mayo del 2009), expresó una profunda preocupación por la baja cantidad de plutonio almacenado, insuficiente para desarrollar todas las misiones de exploración planetaria planeadas en el futuro [81, 91]. Esta circustancia ha llevado a dicha Agencia tomar la decisión de limitar el uso de estos sistemas RPS en algunas misiones de especial interés científico y una recomendación de alta prioridad para que el Congreso de los EEUU apruebe el reestablecimiento de la producción de plutonio-238, -son necesarios cerca de 5 kg de este material radiactivo al año-, para salvaguardar las misiones que requieran dichos sistemas de potencia a partir del año 2018. Por otro lado, la Agencia estadounidense ha estado considerando el uso de fuentes de energía alternativa; como la fisión nuclear a través del ambicioso proyecto Prometheus, para llevar a cabo una misión de exploración en el sistema jovial (JIMO). Finalmente, dicha misión fue desestimada por su elevado coste. Recientemente se han estado desarrollando sistemas que consigan energía a través de los recursos naturales que nos aporta el Sol, mediante paneles solares -poco eficientes para misiones a planetas alejados de la luz solar-. En este contexto, la misión JUNO del programa Nuevas Fronteras de la NASA, cuyo lanzamiento fue realizado con éxito en Agosto de 2011, va a ser la primera misión equipada con paneles solares que sobrevolará Júpiter en el 2015 siguiendo una órbita polar. Anteriormente se habían empleado los antes mencionados RPS para las misiones Pioneer 10,11, Voyager 1,2, Ulysses, Cassini-Huygens y Galileo (todas sobrevuelos excepto Galileo). Dicha misión seguirá una órbita elíptica de alta excentricidad con un periápside muy cercano a Júpiter, y apoápside lejano, evitando que los intensos cinturones de radiación puedan dañar los instrumentos de navegación y científicos. Un tether o amarra electrodinámica es capaz de operar como sistema propulsivo o generador de potencia, pero también puede ser considerado como solución científicotecnológica en misiones espaciales tanto en LEO (Órbita Baja Terrestre) como en planetas exteriores. Siguiendo una perspectiva histórica, durante las misiones terrestres TSS-1 (1992) y TSS-1R (1996) se emplearon amarras estandard con recubrimiento aislante en toda su longitud, aplicando como terminal anódico pasivo un colector esférico para captar electrones. En una geometría alternativa, propuesta por J. R. Sanmartín et al. (1993) [93], se consideró dejar la amarra sin recubrimiento aislante (“bare tether”), y sin colector anódico esférico, de forma que recogiera electrones a lo largo del segmento que resulta polarizado positivo, como si se tratara de una sonda de Langmuir de gran longitud. A diferencia de la amarra estandard, el “bare tether” es capaz de recoger electrones a lo largo de una superficie grande ya que este segmento es de varios kilómetros de longitud. Como el radio de la amarra es del orden de la longitud de Debye y pequeño comparado con el radio de Larmor de los electrones, permite una recolección eficiente de electrones en el régimen OML (Orbital Motion Limited) de sondas de Langmuir. La corriente dada por la teoría OML varía en función del perímetro y la longitud. En el caso de una cinta delgada, el perímetro depende de la anchura, que debe ser suficientemente grande para evitar cortes producidos por debris y micrometeoritos, y suficientemente pequeño para que la amarra funcione en dicho régimen [95]. En el experimento espacial TSS-1R mencionado anteriormente, se identificó una recolección de corriente más elevada que la que predecía el modelo teórico de Parker- Murphy, debido posiblemente a que se utilizaba un colector esférico de radio bastante mayor que la longitud de Debye [79]. En el caso de una amarra “bare”, que recoge electrones a lo largo de gran parte de su longitud, se puede producir un fenómeno conocido como atrapamiento adiabático de electrones (adiabatic electron trapping) [25, 40, 60, 73, 74, 97]. En el caso terrestre (LEO) se da la condición mesotérmica en la que la amarra se mueve con una velocidad muy superior a la velocidad térmica de los iones del ambiente y muy inferior a la velocidad térmica de los electrones. J. Laframboise y L. Parker [57] mostraron que, para una función de distribución quasi-isotrópica, la densidad de electrones debe entonces ser necesariamente inferior a la densidad ambiente. Por otra parte, debido a su flujo hipersónico y a la alta polarización positiva de la amarra, la densidad de los iones es mayor que la densidad ambiente en una vasta región de la parte “ram” del flujo, violando la condición de cuasi-neutralidad,-en una región de dimensión mayor que la longitud de Debye-. La solución a esta paradoja podría basarse en el atrapamiento adiabático de electrones ambiente en órbitas acotadas entorno al tether. ABSTRACT New technological and scientific applications by electrodynamic tethers for planetary missions are analyzed: i) A set of cylindrical, parallel tethers (electric solar sail or e-sail) is considered for an interplanetary mission; ions from the solar wind are repelled by the high potential of the tether, providing momentum to the e-sail. An approximated model of a stationary potential for a high solar wind flow is considered. With the force provided by a negative biased tether, an indirect method for the optimization trajectory of an Earth-to-Jupiter orbit transfer is analyzed. ii) The deployment of a tether from the e-sail allows several scientific applications in Jupiter. iia) It might be used as a source of radiative waves for plasma diagnostics and artificial aurora generator. A conductive tether orbiting in the Jovian magnetosphere produces waves. Wave radiation by a conductor carrying a steady current in both a polar, highly eccentric, low perijove orbit, as in NASA’s Juno mission, and an equatorial low Jovian orbit (LJO) mission below the intense radiation belts, is considered. Both missions will need electric power generation for scientific instruments and communication systems. Tethers generate power more efficiently than solar panels or radioisotope power systems (RPS). The radiation impedance is required to determine the current in the overall tether circuit. In a cold plasma model, radiation occurs mainly in the Alfven and fast magnetosonic modes, exhibiting a large refraction index. The radiation impedance of insulated tethers is determined for both modes and either mission. Unlike the Earth ionospheric case, the low-density, highly magnetized Jovian plasma makes the electron gyrofrequency much larger than the plasma frequency; this substantially modifies the power spectrum for either mode by increasing the Alfven velocity. An estimation of the radiation impedance of bare tethers is also considered. iib) In LJO, a spacecraft orbiting in a slow downward spiral under the radiation belts would allow determining magnetic field structure and atmospheric composition for understanding the formation, evolution, and structure of Jupiter. Additionally, if the cathodic contactor is switched off, a tether floats electrically, allowing e-beam emission that generate auroras. On/off switching produces bias/current pulses and signal emission, which might be used for Jovian plasma diagnostics. In LJO, the ions impacting against the negative-biased tether do produce secondary electrons, which racing down Jupiter’s magnetic field lines, reach the upper atmosphere. The energetic electrons there generate auroral effects. Regions where the tether efficiently should produce secondary electrons are analyzed. iic) Other scientific application suggested in LJO is the in-situ detection of charged grains. Charged grains naturally orbit near Jupiter. High-energy electrons in the Jovian ambient may be modeled by the kappa distribution function. In complex plasma scenarios, where the Jovian high electric field may accelerate charges up superthermal velocities, the use of non-Maxwellian distributions should be considered. In these cases, the distribution tails fit well to a power-law dependence for electrons. Fluctuations of the charged grains for non-Mawellian distribution function are here studied. iii) The present thesis is concluded with the analysis for de-orbiting satellites at end of mission by electrodynamic tethers. A de-orbit tether system must present very small tether-to-satellite mass ratio and small probability of a tether cut by small debris too. The present work shows how to select tape dimensions so as to minimize the product of those two magnitudes. Preliminary results of tape-tether design are here discussed to minimize that function. Results for de-orbiting Cryosat and Envisat are also presented.

Enhanced thermionic currents by non equilibrium electron population of metals

An analytical expression is derived for the electron thermionic current from heated metals by using a non equilibrium, modified Kappa energy distribution for electrons. This isotropic distribution characterizes the long high energy tails in the electron energy spectrum for low values of the index ? and also accounts for the Fermi energy for the metal electrons. The limit for large ? recovers the classical equilibrium Fermi-Dirac distribution. The predicted electron thermionic current for low ? increases between four and five orders of magnitude with respect to the predictions of the equilibrium Richardson-Dushmann current. The observed departures from this classical expression, also recovered for large ?, would correspond to moderate values of this index. The strong increments predicted by the thermionic emission currents suggest that, under appropriate conditions, materials with non equilibrium electron populations would become more efficient electron emitters at low temperatures.

Change of extreme rainfall indexes at Ebro River Basin

The purpose of this work is to provide a description of the heavy rainfall phenomenon on statistical tools from a Spanish region. We want to quantify the effect of the climate change to verify the rapidity of its evolution across the variation of the probability distributions. Our conclusions have special interest for the agrarian insurances, which may make estimates of costs more realistically. In this work, the analysis mainly focuses on: The distribution of consecutive days without rain for each gauge stations and season. We estimate density Kernel functions and Generalized Pareto Distribution (GPD) for a network of station from the Ebro River basin until a threshold value u. We can establish a relation between distributional parameters and regional characteristics. Moreover we analyze especially the tail of the probability distribution. These tails are governed by law of power means that the number of events n can be expressed as the power of another quantity x : n(x) = x? . ? can be estimated as the slope of log-log plot the number of events and the size. The most convenient way to analyze n(x) is using the empirical probability distribution. Pr(X mayor que x) ? x-?. The distribution of rainfall over percentile of order 0.95 from wet days at the seasonal scale and in a yearly scale with the same treatment of tails than in the previous section.

Distinguishing surface effects of calcium ion from pore-occupancy effects in Na+ channels

The effects of calcium ion on the Na+ activation gate were studied in squid giant axons. Saxitoxin (STX) was used to block ion entry into Na+ channels without hindering access to the membrane surface, making it possible to distinguish surface effects of calcium from pore-occupancy effects. In the presence of STX, gating kinetics were measured from gating current (Ig). The kinetic effects of external calcium concentration changes were small when STX was present. In the absence of STX, lowering the calcium concentration (from 100 to 10 mM) slowed the closing of Na+ channels (measured from INa tails) by more than a factor of 2. Surprisingly, the voltage sensitivity of closing kinetics changed with calcium concentration, and it was modified by STX. Voltage sensitivity apparently depends in part on the ability of calcium to enter and block the channels as voltage is driven negative. In external medium with no added calcium, INa tail current initially increases in amplitude severalfold with the relief of calcium block, then progressively slows and gets smaller, as calcium diffuses out of the layers investing the axon. INa tails seen just before the current disappears suggest that closing in the absence of channel block is very slow or does not occur. INa amplitude and kinetics are completely restored when calcium is returned. The results strongly suggest that calcium occupancy is a requirement for channel closing and that nonoccupied channels fold reversibly into a nonfunctional conformation.

Absence of cytokine receptor-dependent specificity in red blood cell differentiation in vivo

Erythropoietin (EPO) is required for red blood cell development, but whether EPO-specific signals directly instruct erythroid differentiation is unknown. We used a dominant system in which constitutively active variants of the EPO receptor were introduced into erythroid progenitors in mice. Chimeric receptors were constructed by replacing the cytoplasmic tail of constitutively active variants of the EPO receptor with tails of diverse cytokine receptors. Receptors linked to granulocyte or platelet production supported complete erythroid development in vitro and in vivo, as did the growth hormone receptor, a nonhematopoietic receptor. Therefore, EPOR-specific signals are not required for terminal differentiation of erythrocytes. Furthermore, we found that cellular context can influence cytokine receptor signaling.

Polyadenylation of stable RNA precursors in vivo

Polyadenylation at the 3′ terminus has long been considered a specific feature of mRNA and a few other unstable RNA species. Here we show that stable RNAs in Escherichia coli can be polyadenylated as well. RNA molecules with poly(A) tails are the major products that accumulate for essentially all stable RNA precursors when RNA maturation is slowed because of the absence of processing exoribonucleases; poly(A) tails vary from one to seven residues in length. The polyadenylation process depends on the presence of poly(A) polymerase I. A stochastic competition between the exoribonucleases and poly(A) polymerase is proposed to explain the accumulation of polyadenylated RNAs. These data indicate that polyadenylation is not unique to mRNA, and its widespread occurrence suggests that it serves a more general function in RNA metabolism.

Exploring the origins of topological frustration: Design of a minimally frustrated model of fragment B of protein A

Topological frustration in an energetically unfrustrated off-lattice model of the helical protein fragment B of protein A from Staphylococcus aureus was investigated. This Gō-type model exhibited thermodynamic and kinetic signatures of a well-designed two-state folder with concurrent collapse and folding transitions and single exponential kinetics at the transition temperature. Topological frustration is determined in the absence of energetic frustration by the distribution of Fersht φ values. Topologically unfrustrated systems present a unimodal distribution sharply peaked at intermediate φ, whereas highly frustrated systems display a bimodal distribution peaked at low and high φ values. The distribution of φ values in protein A was determined both thermodynamically and kinetically. Both methods yielded a unimodal distribution centered at φ = 0.3 with tails extending to low and high φ values, indicating the presence of a small amount of topological frustration. The contacts with high φ values were located in the turn regions between helices I and II and II and III, intimating that these hairpins are in large part required in the transition state. Our results are in good agreement with all-atom simulations of protein A, as well as lattice simulations of a three- letter code 27-mer (which can be compared with a 60-residue helical protein). The relatively broad unimodal distribution of φ values obtained from the all-atom simulations and that from the minimalist model for the same native fold suggest that the structure of the transition state ensemble is determined mostly by the protein topology and not energetic frustration.

Translation initiation factor eIF4G mediates in vitro poly(A) tail-dependent translation

The yeast translation factor eIF4G associates with both the cap-binding protein eIF4E and the poly(A)-binding protein Pab1p. Here we report that the two yeast eIF4G homologs, Tif4631p and Tif4632p, share a conserved Pab1p-binding site. This site is required for Pab1p and poly(A) tails to stimulate the in vitro translation of uncapped polyadenylylated mRNA, and the region encompassing it is required for the cap and the poly(A) tail to synergistically stimulate translation. This region on Tif4631p becomes essential for cell growth when the eIF4E binding site on Tif4631p is mutated. Pab1p mutations also show synthetic lethal interactions with eIF4E mutations. These data suggest that eIF4G mediates poly(A) tail stimulated translation in vitro, and that Pab1p and the domain encompassing the Pab1p-binding site on eIF4G can compensate for partial loss of eIF4E function in vivo.

Direct interaction of Gβγ with a C-terminal Gβγ-binding domain of the Ca2+ channel α1 subunit is responsible for channel inhibition by G protein-coupled receptors

Several classes of voltage-gated Ca2+ channels (VGCCs) are inhibited by G proteins activated by receptors for neurotransmitters and neuromodulatory peptides. Evidence has accumulated to indicate that for non-L-type Ca2+ channels the executing arm of the activated G protein is its βγ dimer (Gβγ). We report below the existence of two Gβγ-binding sites on the A-, B-, and E-type α1 subunits that form non-L-type Ca2+ channels. One, reported previously, is in loop 1 connecting transmembrane domains I and II. The second is located approximately in the middle of the ca. 600-aa-long C-terminal tails. Both Gβγ-binding regions also bind the Ca2+ channel β subunit (CCβ), which, when overexpressed, interferes with inhibition by activated G proteins. Replacement in α1E of loop 1 with that of the G protein-insensitive and Gβγ-binding-negative loop 1 of α1C did not abolish inhibition by G proteins, but the exchange of the α1E C terminus with that of α1C did. This and properties of α1E C-terminal truncations indicated that the Gβγ-binding site mediating the inhibition of Ca2+ channel activity is the one in the C terminus. Binding of Gβγ to this site was inhibited by an α1-binding domain of CCβ, thus providing an explanation for the functional antagonism existing between CCβ and G protein inhibition. The data do not support proposals that Gβγ inhibits α1 function by interacting with the site located in the loop I–II linker. These results define the molecular mechanism by which presynaptic G protein-coupled receptors inhibit neurotransmission.

p23, a major COPI-vesicle membrane protein, constitutively cycles through the early secretory pathway

A novel type I transmembrane protein of COPI-coated vesicles, p23, has been demonstrated to be localized mainly to the Golgi complex. This protein and p24, another member of the p24 family, have been shown to bind coatomer via their short cytoplasmic tails. Here we demonstrate that p23 continuously cycles through the early secretory pathway. The cytoplasmic tail of p23 is shown to act as a functional retrieval signal as it confers endoplasmic reticulum (ER) residence to a CD8–p23 fusion protein. This ER localization is, at least in part, a result of retrieval from post-ER compartments because CD8–p23 fusion proteins receive post-ER modifications. In contrast, the cytoplasmic tail of p24 has been shown not to retrieve a CD8–p24 fusion protein. The coatomer binding motifs FF and KK in the cytoplasmic tail of p23 are reported to influence the steady-state localization of the CD8–p23 fusion protein within the ER–Golgi recycling pathway. It appears that the steady-state Golgi localization of endogenous p23 is maintained by its lumenal domain, as a fusion protein with the lumenal domain of CD8, and the membrane span as well as the cytoplasmic tail of p23 is no longer detected in the Golgi.

Development of an in vitro mRNA decay system for Escherichia coli: Poly(A) polymerase I is necessary to trigger degradation

Using a novel Escherichia coli in vitro decay system in which polysomes are the source of both enzymes and mRNA, we demonstrate a requirement for poly(A) polymerase I (PAP I) in mRNA turnover. The in vitro decay of two different mRNAs (trxA and lpp) is triggered by the addition of ATP only when polysomes are prepared from a strain carrying the wild-type gene for PAP I (pcnB+). The relative decay rates of these two messages are similar in vitro and in vivo. Poly(A) tails are formed on both mRNAs, but no poly(A) tails are detected on the 3′ end of mature 23S rRNA. The size distribution of poly(A) tails generated in vitro, averaging 50 nt in length, is comparable to that previously reported in vivo. PAP I activity is associated exclusively with the polysomes. Exogenously added PAP I does not restore mRNA decay to PAP I− polysomes, suggesting that, in vivo, PAP I may be part of a multiprotein complex. The potential of this in vitro system for analyzing mRNA decay in E. coli is discussed.

Differential binding to HLA-C of p50-activating and p58-inhibitory natural killer cell receptors

Natural killer (NK) cell cytotoxicity is regulated in large part by the expression of NK cell receptors able to bind class I major histocompatibility complex glycoproteins. The receptors associated with recognition of HLA-C allospecificities are the two-domain Ig-like molecules, p50 and p58 proteins, with highly homologous extracellular domains but differing in that they have either an activating or inhibitory function, respectively, depending on the transmembrane domain and cytoplasmic tails that they possess. We have compared the binding to HLA-Cw7 of an inhibitory p58 molecule, NKAT2, the highly homologous activating p50 molecule, clone 49, and a second activating p50 molecule, clone 39, which has homologies to both NKAT1 and NKAT2. NKAT2 binds to HLA-Cw7 with very rapid association and dissociation rates. However, the p50 receptors bind only very weakly, if at all, to HLA-C. The molecular basis of this difference is analyzed, and the functional significance of these observations is discussed.

Structural details of an interaction between cardiolipin and an integral membrane protein

Anionic lipids play a variety of key roles in biomembrane function, including providing the immediate environment for the integral membrane proteins that catalyze photosynthetic and respiratory energy transduction. Little is known about the molecular basis of these lipid–protein interactions. In this study, x-ray crystallography has been used to examine the structural details of an interaction between cardiolipin and the photoreaction center, a key light-driven electron transfer protein complex found in the cytoplasmic membrane of photosynthetic bacteria. X-ray diffraction data collected over the resolution range 30.0–2.1 Å show that binding of the lipid to the protein involves a combination of ionic interactions between the protein and the lipid headgroup and van der Waals interactions between the lipid tails and the electroneutral intramembrane surface of the protein. In the headgroup region, ionic interactions involve polar groups of a number of residues, the protein backbone, and bound water molecules. The lipid tails sit along largely hydrophobic grooves in the irregular surface of the protein. In addition to providing new information on the immediate lipid environment of a key integral membrane protein, this study provides the first, to our knowledge, high-resolution x-ray crystal structure for cardiolipin. The possible significance of this interaction between an integral membrane protein and cardiolipin is considered.