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SUMMARY One of the outstanding problems in modern geodynamics is the development of thermal convection models that are consistent with the present-day flow dynamics in the Earth's mantle, in accord with seismic tomographic images of 3-D Earth structure, and that are also capable of providing a time-dependent evolution of the mantle thermal structure that is as 'realistic' (Earth-like) as possible. A successful realization of this objective would provide a realistic model of 3-D mantle convection that has optimal consistency with a wide suite of seismic, geodynamic and mineral physical constraints on mantle structure and thermodynamic properties. To address this challenge, we have constructed a time-dependent, compressible convection model in 3-D spherical geometry that is consistent with tomography-based instantaneous flow dynamics, using an updated and revised pseudo-spectral numerical method. The novel feature of our numerical solutions is that the equations of conservation of mass and momentum are solved only once in terms of spectral Green's functions. We initially focus on the theory and numerical methods employed to solve the equation of thermal energy conservation using the Green's function solutions for the equation of motion, with special attention placed on the numerical accuracy and stability of the convection solutions. A particular concern is the verification of the global energy balance in the dissipative, compressible-mantle formulation we adopt. Such validation is essential because we then present geodynamically constrained convection solutions over billion-year timescales, starting from present-day seismically constrained thermal images of the mantle. The use of geodynamically constrained spectral Green's functions facilitates the modelling of the dynamic impact on the mantle evolution of: (1) depth-dependent thermal conductivity profiles, (2) extreme variations of viscosity over depth and (3) different surface boundary conditions, in this case mobile surface plates and a rigid surface. The thermal interpretation of seismic tomography models does not provide a radial profile of the horizontally averaged temperature (i.e. the geotherm) in the mantle. One important goal of this study is to obtain a steady-state geotherm with boundary layers which satisfies energy balance of the system and provides the starting point for more realistic numerical simulations of the Earth's evolution. We obtain surface heat flux in the range of Earth-like values : 37 TW for a rigid surface and 44 TW for a surface with tectonic plates coupled to the mantle flow. Also, our convection simulations deliver CMB heat flux that is on the high end of previously estimated values, namely 13 TW and 20 TW, for rigid and plate-like surface boundary conditions, respectively. We finally employ these two end-member surface boundary conditions to explore the very-long-time scale evolution of convection over billion-year time windows. These billion-year-scale simulations will allow us to determine the extent to which a 'memory' of the starting tomography-based thermal structure is preserved and hence to explore the longevity of the structures in the present-day mantle. The two surface boundary conditions, along with the geodynamically inferred radial viscosity profiles, yield steady-state convective flows that are dominated by long wavelengths throughout the lower mantle. The rigid-surface condition yields a spectrum of mantle heterogeneity dominated by spherical harmonic degree 3 and 4, and the plate-like surface condition yields a pattern dominated by degree 1. Our exploration of the time-dependence of the spatial heterogeneity shows that, for both types of surface boundary condition, deep-mantle hot upwellings resolved in the present-day tomography model are durable and stable features. These deeply rooted mantle plumes show remarkable longevity over very long geological time spans, mainly owing to the geodynamically inferred high viscosity in the lower mantle. [ABSTRACT FROM AUTHOR]

Intraplate magmatic provinces found away from plate boundaries provide direct sampling of the composition and heterogeneity of the Earth's mantle. The chemical heterogeneities that have been observed in the mantle are usually attributed to recycling during subduction 1-3 , which allows for the addition of volatiles and incompatible elements into the mantle. Although many intraplate volcanoes sample deep-mantle reservoirs-possibly at the core-mantle boundary 4 -not all intraplate volcanoes are deep-rooted 5 , and reservoirs in other, shallower boundary layers are likely to participate in magma generation. Here we present evidence that suggests Bermuda sampled a previously unknown mantle domain, characterized by silica-undersaturated melts that are substantially enriched in incompatible elements and volatiles, and a unique, extreme isotopic signature. To our knowledge, Bermuda records the most radiogenic 206 Pb/ 204 Pb isotopes that have been documented in an ocean basin (with 206 Pb/ 204 Pb ratios of 19.9-21.7) using high-precision methods. Together with low 207 Pb/ 204 Pb ratios (15.5-15.6) and relatively invariant Sr, Nd, and Hf isotopes, the data suggest that this source must be less than 650 million years old. We therefore interpret the Bermuda source as a previously unknown, transient mantle reservoir that resulted from the recycling and storage of incompatible elements and volatiles 6-8 in the transition zone (between the upper and lower mantle), aided by the fractionation of lead in a mineral that is stable only in this boundary layer, such as K-hollandite 9,10 . We suggest that recent recycling into the transition zone, related to subduction events during the formation of Pangea, is the reason why this reservoir has only been found in the Atlantic Ocean. Our geodynamic models suggest that this boundary layer was sampled by disturbances related to mantle flow. Seismic studies and diamond inclusions 6,7 have shown that recycled materials can be stored in the transition zone 11 . For the first time, to our knowledge, we show geochemical evidence that this storage is key to the generation of extreme isotopic domains that were previously thought to be related only to deep recycling.

Earth's tectonic plates are generally considered to be driven largely by negative buoyancy associated with subduction of oceanic lithosphere. In this context, mid-ocean ridges (MORs) are passive plate boundaries whose divergence accommodates flow driven by subduction of oceanic slabs at trenches. We show that over the past 80 million years (My), the East Pacific Rise (EPR), Earth's dominant MOR, has been characterized by limited ridge-perpendicular migration and persistent, asymmetric ridge accretion that are anomalous relative to other MORs. We reconstruct the subduction-related buoyancy fluxes of plates on either side of the EPR. The general expectation is that greater slab pull should correlate with faster plate motion and faster spreading at the EPR. Moreover, asymmetry in slab pull on either side of the EPR should correlate with either ridge migration or enhanced plate velocity in the direction of greater slab pull. Based on our analysis, none of the expected correlations are evident. This implies that other forces significantly contribute to EPR behavior. We explain these observations using mantle flow calculations based on globally integrated buoyancy distributions that require core-mantle boundary heat flux of up to 20 TW. The time-dependent mantle flow predictions yield a long-lived deep-seated upwelling that has its highest radial velocity under the EPR and is inferred to control its observed kinematics. The mantle-wide upwelling beneath the EPR drives horizontal components of asthenospheric flows beneath the plates that are similarly asymmetric but faster than the overlying surface plates, thereby contributing to plate motions through viscous tractions in the Pacific region.

Sedimentary rocks from Virginia through Florida record marine flooding during the mid-Pliocene. Several wave-cut scarps that at the time of deposition would have been horizontal are now draped over a warped surface with a maximum variation of 60 meters. We modeled dynamic topography by using mantle convection simulations that predict the amplitude and broad spatial distribution of this distortion. The results imply that dynamic topography and, to a lesser extent, glacial isostatic adjustment account for the current architecture of the coastal plain and proximal shelf. This confounds attempts to use regional stratigraphic relations as references for longer-term sea-level determinations. Inferences of Pliocene global sea-level heights or stability of Antarctic ice sheets therefore cannot be deciphered in the absence of an appropriate mantle dynamic reference frame.

The rat auditory cortex is divided anatomically into several areas, but little is known about the functional differences in information processing between these areas. To determine the filter properties of rat posterior auditory field (PAF) neurons, we compared neurophysiological responses to simple tones, frequency modulated (FM) sweeps, and amplitude modulated noise and tones with responses of primary auditory cortex (A1) neurons. PAF neurons have excitatory receptive fields that are on average 65% broader than A1 neurons. The broader receptive fields of PAF neurons result in responses to narrow and broadband inputs that are stronger than A1. In contrast to A1, we found little evidence for an orderly topographic gradient in PAF based on frequency. These neurons exhibit latencies that are twice as long as A1. In response to modulated tones and noise, PAF neurons adapt to repeated stimuli at significantly slower rates. Unlike A1, neurons in PAF rarely exhibit facilitation to rapidly repeated sounds. Neurons in PAF do not exhibit strong selectivity for rate or direction of narrowband one octave FM sweeps. These results indicate that PAF, like nonprimary visual fields, processes sensory information on larger spectral and longer temporal scales than primary cortex.

Classical conditioning paradigms have been shown to cause frequency-specific plasticity in both primary and secondary cortical areas. Previous research demonstrated that repeated pairing of nucleus basalis (NB) stimulation with a tone results in plasticity in primary auditory cortex (A1), mimicking the changes observed after classical conditioning. However, few studies have documented the effects of similar paradigms in secondary cortical areas. The purpose of this study was to quantify plasticity in the posterior auditory field (PAF) of the rat after NB stimulation paired with a high-frequency tone. NB-tone pairing increased the frequency selectivity of PAF sites activated by the paired tone. This frequency-specific receptive field size narrowing led to a reorganization of PAF such that responses to low- and mid-frequency tones were reduced by 40%. Plasticity in A1 was consistent with previous studies -- pairing a high-frequency tone with NB stimulation expanded the high-frequency region of the frequency map. Receptive field sizes did not change, but characteristic frequencies in A1 were shifted after NB-tone pairing. These results demonstrate that experience-dependent plasticity can take different forms in both A1 and secondary auditory cortex.

The brain is constantly adapting to environmental and endogenous changes (including injury) that occur at every stage of life. The mechanisms that regulate neural plasticity have been refined over millions of years. Motivation and sensory experience directly shape the rewiring that makes learning and neurological recovery possible. Guiding neural reorganization in a manner that facilitates recovery of function is a primary goal of neurological rehabilitation. As the rules that govern neural plasticity become better understood, it will be possible to manipulate the sensory and motor experience of patients to induce specific forms of plasticity. This review summarizes our current knowledge regarding factors that regulate cortical plasticity, illustrates specific forms of reorganization induced by control of each factor, and suggests how to exploit these factors for clinical benefit.

Temporal features are important for the identification of natural sounds. Earlier studies have shown that cortical processing of temporal information can be altered by long-term experience with modulated sounds. In a previous study, we observed that environmental enrichment dramatically increased the response of cortical neurons to single tone and noise burst stimuli in both awake and anesthetized rats. Here, we evaluate how enrichment influences temporal information processing in the auditory cortex. We recorded responses to repeated tones and noise bursts in awake rats using epidural evoked potentials and in anesthetized rats using microelectrodes. Enrichment increased the response of cortical neurons to stimuli presented at slow rates and decreased the response to stimuli presented at fast rates relative to controls. Our observation that enrichment substantially increased response strength and forward masking is consistent with earlier reports that long-term potentiation of cortical synapses is associated with increased paired-pulse depression. Enrichment also increased response synchronization at slow rates and decreased synchronization at fast rates. Paired-pulse depression increased within days of environmental enrichment and was restored to normal levels after return to standard housing conditions. These results are relevant to several clinical disorders characterized by abnormal gating of sensory information, including autism, schizophrenia, and dyslexia.

The mammalian auditory system evolved to extract meaningful information from complex acoustic environments. Spectrotemporal selectivity of auditory neurons provides a potential mechanism to represent natural sounds. Experience-dependent plasticity mechanisms can remodel the spectrotemporal selectivity of neurons in primary auditory cortex (A1). Electrical stimulation of the cholinergic nucleus basalis (NB) enables plasticity in A1 that parallels natural learning and is specific to acoustic features associated with NB activity. In this study, we used NB stimulation to explore how cortical networks reorganize after experience with frequency-modulated (FM) sweeps, and how background stimuli contribute to spectrotemporal plasticity in rat auditory cortex. Pairing an 8-4 kHz FM sweep with NB stimulation 300 times per day for 20 days decreased tone thresholds, frequency selectivity, and response latency of A1 neurons in the region of the tonotopic map activated by the sound. In an attempt to modify neuronal response properties across all of A1 the same NB activation was paired in a second group of rats with five downward FM sweeps, each spanning a different octave. No changes in FM selectivity or receptive field (RF) structure were observed when the neural activation was distributed across the cortical surface. However, the addition of unpaired background sweeps of different rates or direction was sufficient to alter RF characteristics across the tonotopic map in a third group of rats. These results extend earlier observations that cortical neurons can develop stimulus specific plasticity and indicate that background conditions can strongly influence cortical plasticity.

Correlation-based synaptic plasticity provides a potential cellular mechanism for learning and memory. Studies in the visual and somatosensory systems have shown that behavioral and surgical manipulation of sensory inputs leads to changes in cortical organization that are consistent with the operation of these learning rules. In this study, we examine how the organization of primary auditory cortex (A1) is altered by tones designed to decrease the average input correlation across the frequency map. After one month of separately pairing nucleus basalis stimulation with 2 and 14 kHz tones, a greater proportion of A1 neurons responded to frequencies below 2 kHz and above 14 kHz. Despite the expanded representation of these tones, cortical excitability was specifically reduced in the high and low frequency regions of A1, as evidenced by increased neural thresholds and decreased response strength. In contrast, in the frequency region between the two paired tones, driven rates were unaffected and spontaneous firing rate was increased. Neural response latencies were increased across the frequency map when nucleus basalis stimulation was associated with asynchronous activation of the high and low frequency regions of A1. This set of changes did not occur when pulsed noise bursts were paired with nucleus basalis stimulation. These results are consistent with earlier observations that sensory input statistics can shape cortical map organization and spike timing.

Over the last 50 yr, environmental enrichment has been shown to generate more than a dozen changes in brain anatomy. The consequences of these physical changes on information processing have not been well studied. In this study, rats were housed in enriched or standard conditions either prior to or after reaching sexual maturity. Evoked potentials from awake rats and extracellular recordings from anesthetized rats were used to document responses of auditory cortex neurons. This report details several significant, new findings about the influence of housing conditions on the responses of rat auditory cortex neurons. First, enrichment dramatically increases the strength of auditory cortex responses. Tone-evoked potentials of enriched rats, for example, were more than twice the amplitude of rats raised in standard laboratory conditions. Second, cortical responses of both young and adult animals benefit from exposure to an enriched environment and are degraded by exposure to an impoverished environment. Third, housing condition resulted in rapid remodeling of cortical responses in <2 wk. Fourth, recordings made under anesthesia indicate that enrichment increases the number of neurons activated by any sound. This finding shows that the evoked potential plasticity documented in awake rats was not due to differences in behavioral state. Finally, enrichment made primary auditory cortex (A1) neurons more sensitive to quiet sounds, more selective for tone frequency, and altered their response latencies. These experiments provide the first evidence of physiologic changes in auditory cortex processing resulting from generalized environmental enrichment.

Sensory experience alters the functional organization of cortical networks. Previous studies using behavioral training motivated by aversive or rewarding stimuli have demonstrated that cortical plasticity is specific to salient inputs in the sensory environment. Sensory experience associated with electrical activation of the basal forebrain (BasF) generates similar input specific plasticity. By directly engaging plasticity mechanisms and avoiding extensive behavioral training, BasF stimulation makes it possible to efficiently explore how specific sensory features contribute to cortical plasticity. This review summarizes our observations that cortical networks employ a variety of strategies to improve the representation of the sensory environment. Different combinations of receptive-field, temporal, and spectrotemporal plasticity were generated in primary auditory cortex neurons depending on the pitch, modulation rate, and order of sounds paired with BasF stimulation. Simple tones led to map expansion, while modulated tones altered the maximum cortical following rate. Exposure to complex acoustic sequences led to the development of combination-sensitive responses. This remodeling of cortical response characteristics may reflect changes in intrinsic cellular mechanisms, synaptic efficacy, and local neuronal connectivity. The intricate relationship between the pattern of sensory activation and cortical plasticity suggests that network-level rules alter the functional organization of the cortex to generate the most behaviorally useful representation of the sensory environment.