Practical Characterization of Input-Parallel-Connected Converters with a Common Input Filter

The paper presents a method to analyze robust stability and transient performance of a distributed power system consisting of commercial converter modules interconnected through a common input filter. The method is based on the use of four transfer functions, which are measurable from the converter input and output terminals. It is shown that these parameters provide important information on the power module sensitivity to the interactions caused by the external impedances. Practical characterization for the described system structure is performed introducing special transfer functions utilized for the interactions assessment. Experimental results are provided to support the presented analysis procedure.

Multiple Input-Output Bidirectional Solid State Transformer Based on a Series Resonant Converter

Las fuentes de alimentación de modo conmutado (SMPS en sus siglas en inglés) se utilizan ampliamente en una gran variedad de aplicaciones. La tarea más difícil para los diseñadores de SMPS consiste en lograr simultáneamente la operación del convertidor con alto rendimiento y alta densidad de energía. El tamaño y el peso de un convertidor de potencia está dominado por los componentes pasivos, ya que estos elementos son normalmente más grandes y más pesados que otros elementos en el circuito. Para una potencia de salida dada, la cantidad de energía almacenada en el convertidor que ha de ser entregada a la carga en cada ciclo de conmutación, es inversamente proporcional a la frecuencia de conmutación del convertidor. Por lo tanto, el aumento de la frecuencia de conmutación se considera un medio para lograr soluciones más compactas con los niveles de densidad de potencia más altos. La importancia de investigar en el rango de alta frecuencia de conmutación radica en todos los beneficios que se pueden lograr: además de la reducción en el tamaño de los componentes pasivos, el aumento de la frecuencia de conmutación puede mejorar significativamente prestaciones dinámicas de convertidores de potencia. Almacenamiento de energía pequeña y el período de conmutación corto conducen a una respuesta transitoria del convertidor más rápida en presencia de las variaciones de la tensión de entrada o de la carga. Las limitaciones más importantes del incremento de la frecuencia de conmutación se relacionan con mayores pérdidas del núcleo magnético convencional, así como las pérdidas de los devanados debido a los efectos pelicular y proximidad. También, un problema potencial es el aumento de los efectos de los elementos parásitos de los componentes magnéticos - inductancia de dispersión y la capacidad entre los devanados - que causan pérdidas adicionales debido a las corrientes no deseadas. Otro factor limitante supone el incremento de las pérdidas de conmutación y el aumento de la influencia de los elementos parásitos (pistas de circuitos impresos, interconexiones y empaquetado) en el comportamiento del circuito. El uso de topologías resonantes puede abordar estos problemas mediante el uso de las técnicas de conmutaciones suaves para reducir las pérdidas de conmutación incorporando los parásitos en los elementos del circuito. Sin embargo, las mejoras de rendimiento se reducen significativamente debido a las corrientes circulantes cuando el convertidor opera fuera de las condiciones de funcionamiento nominales. A medida que la tensión de entrada o la carga cambian las corrientes circulantes incrementan en comparación con aquellos en condiciones de funcionamiento nominales. Se pueden obtener muchos beneficios potenciales de la operación de convertidores resonantes a más alta frecuencia si se emplean en aplicaciones con condiciones de tensión de entrada favorables como las que se encuentran en las arquitecturas de potencia distribuidas. La regulación de la carga y en particular la regulación de la tensión de entrada reducen tanto la densidad de potencia del convertidor como el rendimiento. Debido a la relativamente constante tensión de bus que se encuentra en arquitecturas de potencia distribuidas los convertidores resonantes son adecuados para el uso en convertidores de tipo bus (transformadores cc/cc de estado sólido). En el mercado ya están disponibles productos comerciales de transformadores cc/cc de dos puertos que tienen muy alta densidad de potencia y alto rendimiento se basan en convertidor resonante serie que opera justo en la frecuencia de resonancia y en el orden de los megahercios. Sin embargo, las mejoras futuras en el rendimiento de las arquitecturas de potencia se esperan que vengan del uso de dos o más buses de distribución de baja tensión en vez de una sola. Teniendo eso en cuenta, el objetivo principal de esta tesis es aplicar el concepto del convertidor resonante serie que funciona en su punto óptimo en un nuevo transformador cc/cc bidireccional de puertos múltiples para atender las necesidades futuras de las arquitecturas de potencia. El nuevo transformador cc/cc bidireccional de puertos múltiples se basa en la topología de convertidor resonante serie y reduce a sólo uno el número de componentes magnéticos. Conmutaciones suaves de los interruptores hacen que sea posible la operación en las altas frecuencias de conmutación para alcanzar altas densidades de potencia. Los problemas posibles con respecto a inductancias parásitas se eliminan, ya que se absorben en los Resumen elementos del circuito. El convertidor se caracteriza con una muy buena regulación de la carga propia y cruzada debido a sus pequeñas impedancias de salida intrínsecas. El transformador cc/cc de puertos múltiples opera a una frecuencia de conmutación fija y sin regulación de la tensión de entrada. En esta tesis se analiza de forma teórica y en profundidad el funcionamiento y el diseño de la topología y del transformador, modelándolos en detalle para poder optimizar su diseño. Los resultados experimentales obtenidos se corresponden con gran exactitud a aquellos proporcionados por los modelos. El efecto de los elementos parásitos son críticos y afectan a diferentes aspectos del convertidor, regulación de la tensión de salida, pérdidas de conducción, regulación cruzada, etc. También se obtienen los criterios de diseño para seleccionar los valores de los condensadores de resonancia para lograr diferentes objetivos de diseño, tales como pérdidas de conducción mínimas, la eliminación de la regulación cruzada o conmutación en apagado con corriente cero en plena carga de todos los puentes secundarios. Las conmutaciones en encendido con tensión cero en todos los interruptores se consiguen ajustando el entrehierro para obtener una inductancia magnetizante finita en el transformador. Se propone, además, un cambio en los señales de disparo para conseguir que la operación con conmutaciones en apagado con corriente cero de todos los puentes secundarios sea independiente de la variación de la carga y de las tolerancias de los condensadores resonantes. La viabilidad de la topología propuesta se verifica a través una extensa tarea de simulación y el trabajo experimental. La optimización del diseño del transformador de alta frecuencia también se aborda en este trabajo, ya que es el componente más voluminoso en el convertidor. El impacto de de la duración del tiempo muerto y el tamaño del entrehierro en el rendimiento del convertidor se analizan en un ejemplo de diseño de transformador cc/cc de tres puertos y cientos de vatios de potencia. En la parte final de esta investigación se considera la implementación y el análisis de las prestaciones de un transformador cc/cc de cuatro puertos para una aplicación de muy baja tensión y de decenas de vatios de potencia, y sin requisitos de aislamiento. Abstract Recently, switch mode power supplies (SMPS) have been used in a great variety of applications. The most challenging issue for designers of SMPS is to achieve simultaneously high efficiency operation at high power density. The size and weight of a power converter is dominated by the passive components since these elements are normally larger and heavier than other elements in the circuit. If the output power is constant, the stored amount of energy in the converter which is to be delivered to the load in each switching cycle is inversely proportional to the converter’s switching frequency. Therefore, increasing the switching frequency is considered a mean to achieve more compact solutions at higher power density levels. The importance of investigation in high switching frequency range comes from all the benefits that can be achieved. Besides the reduction in size of passive components, increasing switching frequency can significantly improve dynamic performances of power converters. Small energy storage and short switching period lead to faster transient response of the converter against the input voltage and load variations. The most important limitations for pushing up the switching frequency are related to increased conventional magnetic core loss as well as the winding loss due to the skin and proximity effect. A potential problem is also increased magnetic parasitics – leakage inductance and capacitance between the windings – that cause additional loss due to unwanted currents. Higher switching loss and the increased influence of printed circuit boards, interconnections and packaging on circuit behavior is another limiting factor. Resonant power conversion can address these problems by using soft switching techniques to reduce switching loss incorporating the parasitics into the circuit elements. However the performance gains are significantly reduced due to the circulating currents when the converter operates out of the nominal operating conditions. As the input voltage or the load change the circulating currents become higher comparing to those ones at nominal operating conditions. Multiple Input-Output Many potential gains from operating resonant converters at higher switching frequency can be obtained if they are employed in applications with favorable input voltage conditions such as those found in distributed power architectures. Load and particularly input voltage regulation reduce a converter’s power density and efficiency. Due to a relatively constant bus voltage in distributed power architectures the resonant converters are suitable for bus voltage conversion (dc/dc or solid state transformation). Unregulated two port dc/dc transformer products achieving very high power density and efficiency figures are based on series resonant converter operating just at the resonant frequency and operating in the megahertz range are already available in the market. However, further efficiency improvements of power architectures are expected to come from using two or more separate low voltage distribution buses instead of a single one. The principal objective of this dissertation is to implement the concept of the series resonant converter operating at its optimum point into a novel bidirectional multiple port dc/dc transformer to address the future needs of power architectures. The new multiple port dc/dc transformer is based on a series resonant converter topology and reduces to only one the number of magnetic components. Soft switching commutations make possible high switching frequencies to be adopted and high power densities to be achieved. Possible problems regarding stray inductances are eliminated since they are absorbed into the circuit elements. The converter features very good inherent load and cross regulation due to the small output impedances. The proposed multiple port dc/dc transformer operates at fixed switching frequency without line regulation. Extensive theoretical analysis of the topology and modeling in details are provided in order to compare with the experimental results. The relationships that show how the output voltage regulation and conduction losses are affected by the circuit parasitics are derived. The methods to select the resonant capacitor values to achieve different design goals such as minimum conduction losses, elimination of cross regulation or ZCS operation at full load of all the secondary side bridges are discussed. ZVS turn-on of all the switches is achieved by relying on the finite magnetizing inductance of the Abstract transformer. A change of the driving pattern is proposed to achieve ZCS operation of all the secondary side bridges independent on load variations or resonant capacitor tolerances. The feasibility of the proposed topology is verified through extensive simulation and experimental work. The optimization of the high frequency transformer design is also addressed in this work since it is the most bulky component in the converter. The impact of dead time interval and the gap size on the overall converter efficiency is analyzed on the design example of the three port dc/dc transformer of several hundreds of watts of the output power for high voltage applications. The final part of this research considers the implementation and performance analysis of the four port dc/dc transformer in a low voltage application of tens of watts of the output power and without isolation requirements.

Modeling impact assessment of transport policies through a multiregional input-output integrated approach

El objetivo de esta investigación es desarrollar una metodología para estimar los potenciales impactos económicos y de transporte generados por la aplicación de políticas en el sector transporte. Los departamentos de transporte y otras instituciones gubernamentales relacionadas se encuentran interesadas en estos análisis debido a que son presentados comúnmente de forma errónea por la insuficiencia de datos o por la falta de metodologías adecuadas. La presente investigación tiene por objeto llenar este vacío haciendo un análisis exhaustivo de las técnicas disponibles que coincidan con ese propósito. Se ha realizado un análisis que ha identificado las diferencias cuando son aplicados para la valoración de los beneficios para el usuario o para otros efectos como aspectos sociales. Como resultado de ello, esta investigación ofrece un enfoque integrado que incluye un modelo Input-Output de múltiples regiones basado en la utilidad aleatoria (RUBMRIO), y un modelo de red de transporte por carretera. Este modelo permite la reproducción con mayor detalle y realismo del transporte de mercancías que por medio de su estructura sectorial identifica los vínculos de las compras y ventas inter-industriales dentro de un país utilizando los servicios del transporte de mercancías. Por esta razón, el modelo integrado es aplicable a diversas políticas de transporte. En efecto, el enfoque se ha aplicado para estudiar los efectos macroeconómicos regionales de la implementación de dos políticas diferentes en el sistema de transporte de mercancías de España, tales como la tarificación basada en la distancia recorrida por vehículo-kilómetro (€/km) aplicada a los vehículos del transporte de mercancías, y para la introducción de vehículos más largos y pesados de mercancías en la red de carreteras de España. El enfoque metodológico se ha evaluado caso por caso teniendo en cuenta una selección de la red de carreteras que unen las capitales de las regiones españolas. También se ha tenido en cuenta una dimensión económica a través de una tabla Input-Output de múltiples regiones (MRIO) y la base de datos de conteo de tráfico existente para realizar la validación del modelo. El enfoque integrado reproduce las condiciones de comercio observadas entre las regiones usando el sistema de transporte de mercancías por carretera, y que permite por comparación con los escenarios de políticas, determinar las contribuciones a los cambios distributivos y generativos. Así pues, el análisis estima los impactos económicos en cualquier región considerando los cambios en el Producto Interno Bruto (PIB) y el empleo. El enfoque identifica los cambios en el sistema de transporte a través de todos los caminos de la red de transporte a través de las medidas de efectividad (MOEs). Los resultados presentados en esta investigación proporcionan evidencia sustancial de que en la evaluación de las políticas de transporte, es necesario establecer un vínculo entre la estructura económica de las regiones y de los servicios de transporte. Los análisis muestran que para la mayoría de las regiones del país, los cambios son evidentes para el PIB y el empleo, ya que el comercio se fomenta o se inhibe. El enfoque muestra cómo el tráfico se desvía en ambas políticas, y también determina detalles de las emisiones de contaminantes en los dos escenarios. Además, las políticas de fijación de precios o de regulación de los sistemas de transporte de mercancías por carretera dirigidas a los productores y consumidores en las regiones promoverán transformaciones regionales afectando todo el país, y esto conduce a conclusiones diferentes. Así mismo, este enfoque integrado podría ser útil para evaluar otras políticas y otros países en todo el mundo. The purpose of this research is to develop a methodological approach aimed at assessing the potential economic and transportation impacts of transport policies. Transportation departments and other related government parties are interested in such analysis because it is commonly misrepresented for the insufficiency of data and suitable methodologies available. This research is directed at filling this gap by making a comprehensive analysis of the available techniques that match with that purpose. The differences when they are applied for the valuation of user benefits or for other impacts as social matters have been identified. As a result, this research presents an integrated approach which includes both a random utility-based multiregional Input-Output model (RUBMRIO), and a road transport network model. This model accounts for freight transport with more detail and realism because its commodity-based structure traces the linkages of inter-industry purchases and sales that use freight services within a given country. For this reason, the integrated model is applicable to various transport policies. In fact, the approach is applied to study the regional macroeconomic effects of implementing two different policies in the freight transport system of Spain, such as a distance-based charge in vehicle-kilometer (€/km) for Heavy Goods Vehicles (HGVs), and the introduction of Longer and Heavier Vehicles (LHVs) in the road network of Spain. The methodological approach has been evaluated on a case by case basis considering a selected road network of highways linking the capitals of the Spanish regions. It has also considered an economic dimension through a Multiregional Input Output Table (MRIO) and the existing traffic count database used in the model validation. The integrated approach replicates observed conditions of trade among regions using road freight transport systems that determine contributions to distributional and generative changes by comparison with policy scenarios. Therefore, the model estimates economic impacts in any given area by considering changes in Gross Domestic Product (GDP), employment (jobs), and in the transportation system across all paths of the transport network considering Measures of effectiveness (MOEs). The results presented in this research provide substantive evidence that in the assessment of transport policies it is necessary to establish a link between the economic structure of regions and the transportation services. The analysis shows that for most regions in the country, GDP and employment changes are noticeable when trade is encouraged or discouraged. This approach shows how traffic is diverted in both policies, and also provides details of the pollutant emissions in both scenarios. Furthermore, policies, such as pricing or regulation of road freight transportation systems, directed to producers and consumers in regions will promote different regional transformations across the country, and this lead to different conclusions. In addition, this integrated approach could be useful to assess other policies and countries worldwide.

Even-sowing pattern strategies for a low-input agricultural system in forage maize

The objective of this study was to verify the effectiveness of new patterns of sowing and to achieve a low-input organic system in two different environments (northern and southern Europe). The study was motivated by the hypothesis that more even sowing patterns (triangular and square) would significantly enhance the growth and yield of forage maize under widely varying conditions, compared with traditional mechanised rectangular seed patterns. An experiment was conducted in Madrid and duplicated in Copenhagen during 2010. A random block design was used with a 2 × 2 factorial arrangement based on two seed-sowing patterns: traditional (rectangular) and new (even) and two weed-management conditions (herbicide use and a low-input system). In both weed-management conditions and locations, the production of aerial maize biomass was greater for the new square seed patterns. In addition, the new pattern showed a greater effectiveness in the control of weeds, both at the initial crop stages (36 and 33% fewer weeds m-2 at the 4- and 8-leaf stages, respectively, in the Copenhagen field experiment) and at the final stage. The final weed biomass for the new pattern was 568 kg ha-1 lower for the Copenhagen experiment and 277 kg ha-1 lower in Madrid field experiments. In the light of these results, the new pattern could potentially reduce the use of herbicides. The results of the experiments support the hypothesis formulated at the beginning of this study that even-sowing patterns would be relatively favourable for the growth and yield of the maize crop. In the near future, new machinery could be used to achieve new seed patterns for the optimisation of biomass yield under low-input systems. This approach is effective because it promotes natural crop-weed competition.

Design energy input spectra for high seismicity regions based on Turkish registers

This work proposes design energy spectra in terms of an equivalent velocity, intended for regions with design peak acceleration 0.3 g or higher. These spectra were derived through linear and nonlinear dynamic analyses on a number of selected Turkish strong ground motion records. In the long and mid period ranges the analyses are linear, given the relative insensitivity of the spectra to structural parameters other than the fundamental period; conversely, in the short period range, the spectra are more sensitive to the structural parameters and, hence, nonlinear analyses are required. The selected records are classified in eight groups with respect to soil type (stiff or soft soil), the severity of the earthquake in terms of surface magnitude Ms(Ms≤ 5.5 and Ms> 5.5) and the relevance of the near-source effects (impulsive or vibratory). For each of these groups, median and characteristic spectra are proposed; such levels would respectively correspond to 50 and 95 % percentiles. These spectra have an initial linear growing branch in the short period range, a horizontal branch in the mid period range and a descending branch in the long period range. Empirical criteria for estimating the hysteretic energy from the input energy are suggested. The proposed design spectra are compared with those obtained from other studies.

Methodology for assessing regional economic impacts of charging HGVs in Spain: an integrated approach through a random utility based multiregional input-output and a road transportation network model

An integrated approach composed of a random utility-based multiregional input-output model and a road transport network model was developed for evaluating the application of a fee to heavy-goods vehicles (HGVs) in Spain. For this purpose, a distance-based charge scenario (in euros per vehicle kilometer) for HGVs was evaluated for a selected motorway network in Spain. Although the aim of this charging policy was to increase the efficiency of transport, the approach strongly identified direct and indirect impacts on the regional economy. Estimates of the magnitude and extent of indirect effects on aggregated macroeconomic indicators (employment and gross domestic product) are provided. The macroeconomic effects of the charging policy were found to be positive for some regions and negative for other regions.

A random utility based multiregional input-output approach to estimate the impact of a new highway on the regional distribution of jobs

This paper addresses the economic impact assessment of the construction of a new road on the regional distribution of jobs. The paper summarizes different existing model approaches considered to assess economic impacts through a literature review. Afterwards, we present the development of a comprehensive approach for analyzing the interaction of new transport infrastructure and the economic impact through an integrated model. This model has been applied to the construction of the motorway A-40 in Spain (497 Km.) which runs across three regions without passing though Madrid City. This may in turn lead to the relocation of labor and capital due to the improvement of accessibility of markets or inputs. The result suggests the existence of direct and indirect effects in other regions derived from the improvement of the transportation infrastructure, and confirms the relevance of road freight transport in some regions. We found that the changes in regional employment are substantial for some regions (increasing or decreasing jobs), but a t the same time negligible in other regions. As a result,the approach provides broad guidance to national governments and other transport-related parties about the impacts of this transport policy.

Design and Multi-Objective Optimization of EMI Input Filters

An EMI filter design procedure for power converters is proposed. Based on a given noise spectrum, information about the converter noise source impedance and design constraints, the design space of the input filter is defined. The design is based on component databases and detailed models of the filter components, including high frequency parasitics, losses, weight, volume, etc.. The design space is mapped onto a performance space in which different filter implementations are evaluated and compared. A multi-objective optimization approach is used to obtain optimal designs w.r.t. a given performance function.

Differential roles of Ca2+/calmodulin-dependent kinases in posttetanic potentiation at input selective glutamatergic pathways

The electrosensory lateral line lobe (ELL) of the electric fish Apteronotus leptorhynchus is a layered medullary region receiving electroreceptor input that terminates on basal dendrites of interneurons and projection (pyramidal) cells. The molecular layer of the ELL contains two distinct glutamatergic feedback pathways that terminate on the proximal (ventral molecular layer, VML) and distal (dorsal molecular layer) apical dendrites of pyramidal cells. Western blot analysis with an antibody directed against mammalian Ca2+/calmodulin-dependent kinase 2, α subunit (CaMK2α) recognized a protein of identical size in the brain of A. leptorhynchus. Immunohistochemistry demonstrated that CaMK2 α expression in the ELL was restricted to fibers and terminals in the VML. Posttetanic potentiation (PTP) could be readily elicited in pyramidal cells by stimulation of either VML or DML in brain slices of the ELL. PTP in the VML was blocked by extracellular application of a CaMK2 antagonist (KN62) while intracellular application of KN62 or a CaMK2 inhibitory peptide had no effect, consistent with the presynaptic localization of CaMK2 α in VML. PTP in the dorsal molecular layer was not affected by extracellular application of KN62. Anti-Hebbian plasticity has also been demonstrated in the VML, but was not affected by KN62. These results demonstrate that, while PTP can occur independent of CaMK2, it is, in some synapses, dependent on this kinase.

The roles of the two proton input channels in cytochrome c oxidase from Rhodobacter sphaeroides probed by the effects of site-directed mutations on time-resolved electrogenic intraprotein proton transfer

The crystal structures of cytochrome c oxidase from both bovine and Paracoccus denitrificans reveal two putative proton input channels that connect the heme-copper center, where dioxygen is reduced, to the internal aqueous phase. In this work we have examined the role of these two channels, looking at the effects of site-directed mutations of residues observed in each of the channels of the cytochrome c oxidase from Rhodobacter sphaeroides. A photoelectric technique was used to monitor the time-resolved electrogenic proton transfer steps associated with the photo-induced reduction of the ferryl-oxo form of heme a3 (Fe4+ = O2−) to the oxidized form (Fe3+OH−). This redox step requires the delivery of a “chemical” H+ to protonate the reduced oxygen atom and is also coupled to proton pumping. It is found that mutations in the K channel (K362M and T359A) have virtually no effect on the ferryl-oxo-to-oxidized (F-to-Ox) transition, although steady-state turnover is severely limited. In contrast, electrogenic proton transfer at this step is strongly suppressed by mutations in the D channel. The results strongly suggest that the functional roles of the two channels are not the separate delivery of chemical or pumped protons, as proposed recently [Iwata, S., Ostermeier, C., Ludwig, B. & Michel, H. (1995) Nature (London) 376, 660–669]. The D channel is likely to be involved in the uptake of both “chemical” and “pumped” protons in the F-to-Ox transition, whereas the K channel is probably idle at this partial reaction and is likely to be used for loading the enzyme with protons at some earlier steps of the catalytic cycle. This conclusion agrees with different redox states of heme a3 in the K362M and E286Q mutants under aerobic steady-state turnover conditions.

Impact of Spatial Soil and Climate Input Data Aggregation on Regional Yield Simulations

This work was financially supported by the German Federal Ministry of Food and Agriculture (BMEL) through the Federal Office for Agriculture and Food (BLE), (2851ERA01J). FT and RPR were supported by FACCE MACSUR (3200009600) through the Finnish Ministry of Agriculture and Forestry (MMM). EC, HE and EL were supported by The Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (220-2007-1218) and by the strategic funding ‘Soil-Water-Landscape’ from the faculty of Natural Resources and Agricultural Sciences (Swedish University of Agricultural Sciences) and thank professor P-E Jansson (Royal Institute of Technology, Stockholm) for support. JC, HR and DW thank the INRA ACCAF metaprogramm for funding and Eric Casellas from UR MIAT INRA for support. CB was funded by the Helmholtz project “REKLIM—Regional Climate Change”. CK was funded by the HGF Alliance “Remote Sensing and Earth System Dynamics” (EDA). FH was funded by the German Research Foundation (Deutsche Forschungsgemeinschaft, DFG) under the Grant FOR1695. FE and SS acknowledge support by the German Science Foundation (project EW 119/5-1). HH, GZ, SS, TG and FE thank Andreas Enders and Gunther Krauss (INRES, University of Bonn) for support. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Control of time-dependent biological processes by temporally patterned input

Temporal patterning of biological variables, in the form of oscillations and rhythms on many time scales, is ubiquitous. Altering the temporal pattern of an input variable greatly affects the output of many biological processes. We develop here a conceptual framework for a quantitative understanding of such pattern dependence, focusing particularly on nonlinear, saturable, time-dependent processes that abound in biophysics, biochemistry, and physiology. We show theoretically that pattern dependence is governed by the nonlinearity of the input–output transformation as well as its time constant. As a result, only patterns on certain time scales permit the expression of pattern dependence, and processes with different time constants can respond preferentially to different patterns. This has implications for temporal coding and decoding, and allows differential control of processes through pattern. We show how pattern dependence can be quantitatively predicted using only information from steady, unpatterned input. To apply our ideas, we analyze, in an experimental example, how muscle contraction depends on the pattern of motorneuron firing.

Decoding temporally encoded sensory input by cortical oscillations and thalamic phase comparators

The temporally encoded information obtained by vibrissal touch could be decoded “passively,” involving only input-driven elements, or “actively,” utilizing intrinsically driven oscillators. A previous study suggested that the trigeminal somatosensory system of rats does not obey the bottom-up order of activation predicted by passive decoding. Thus, we have tested whether this system obeys the predictions of active decoding. We have studied cortical single units in the somatosensory cortices of anesthetized rats and guinea pigs and found that about a quarter of them exhibit clear spontaneous oscillations, many of them around whisking frequencies (≈10 Hz). The frequencies of these oscillations could be controlled locally by glutamate. These oscillations could be forced to track the frequency of induced rhythmic whisker movements at a stable, frequency-dependent, phase difference. During these stimulations, the response intensities of multiunits at the thalamic recipient layers of the cortex decreased, and their latencies increased, with increasing input frequency. These observations are consistent with thalamocortical loops implementing phase-locked loops, circuits that are most efficient in decoding temporally encoded information like that obtained by active vibrissal touch. According to this model, and consistent with our results, populations of thalamic “relay” neurons function as phase “comparators” that compare cortical timing expectations with the actual input timing and represent the difference by their population output rate.

The circadian clock controls the expression pattern of the circadian input photoreceptor, phytochrome B

Developmental and physiological responses are regulated by light throughout the entire life cycle of higher plants. To sense changes in the light environment, plants have developed various photoreceptors, including the red/far-red light-absorbing phytochromes and blue light-absorbing cryptochromes. A wide variety of physiological responses, including most light responses, also are modulated by circadian rhythms that are generated by an endogenous oscillator, the circadian clock. To provide information on local time, circadian clocks are synchronized and entrained by environmental time cues, of which light is among the most important. Light-driven entrainment of the Arabidopsis circadian clock has been shown to be mediated by phytochrome A (phyA), phytochrome B (phyB), and cryptochromes 1 and 2, thus affirming the roles of these photoreceptors as input regulators to the plant circadian clock. Here we show that the expression of PHYB∷LUC reporter genes containing the promoter and 5′ untranslated region of the tobacco NtPHYB1 or Arabidopsis AtPHYB genes fused to the luciferase (LUC) gene exhibit robust circadian oscillations in transgenic plants. We demonstrate that the abundance of PHYB RNA retains this circadian regulation and use a PHYB∷Luc fusion protein to show that the rate of PHYB synthesis is also rhythmic. The abundance of bulk PHYB protein, however, exhibits only weak circadian rhythmicity, if any. These data suggest that photoreceptor gene expression patterns may be significant in the daily regulation of plant physiology and indicate an unexpectedly intimate relationship between the components of the input pathway and the putative circadian clock mechanism in higher plants.