Deep Equatorial Pacific Ocean Oxygenation and Atmospheric CO2 Over The Last Ice Age.

Ventilation of carbon stored in the deep ocean is thought to play an important role in atmospheric CO2 increases associated with Pleistocene deglaciations. The presence of this respired carbon has been recorded by an array of paleoceanographic proxies from various locations across the global ocean. Here we present a new sediment core from the Eastern Equatorial Pacific (EEP) Ocean spanning the last 180,000 years and reconstruct high-resolution 230Th-derived fluxes of 232Th and excess barium, along with redox-sensitive uranium concentrations to examine past variations in dust delivery, export productivity, and bottom-water oxygenation, respectively. Our bottom-water oxygenation record is compared to other similar high-resolution records from across the Pacific and in the Southern Ocean. We suggest that the deep Pacific is a site of respired carbon storage associated with periods of decreased global atmospheric CO2 concentration during the LGM, confirming the conclusions from a wealth of previous studies. However, our study is the first to show a similar relationship beyond the last glacial, extending to at least 70,000 years.

Deglacial Tropical Atlantic subsurface warming links ocean circulation variability to the West African Monsoon.

Multiple lines of evidence show that cold stadials in the North Atlantic were accompanied by both reductions in Atlantic Meridional Overturning Circulation (AMOC) and collapses of the West African Monsoon (WAM). Although records of terrestrial change identify abrupt WAM variability across the deglaciation, few studies show how ocean temperatures evolved across the deglaciation. To identify the mechanism linking AMOC to the WAM, we generated a new record of subsurface temperature variability over the last 21 kyr based on Mg/Ca ratios in a sub-thermocline dwelling planktonic foraminifera in an Eastern Equatorial Atlantic (EEA) sediment core from the Niger Delta. Our subsurface temperature record shows abrupt subsurface warming during both the Younger Dryas (YD) and Heinrich Event 1. We also conducted a new transient coupled ocean-atmosphere model simulation across the YD that better resolves the western boundary current dynamics and find a strong negative correlation between AMOC strength and EEA subsurface temperatures caused by changes in ocean circulation and rainfall responses that are consistent with the observed WAM change. Our combined proxy and modeling results provide the first evidence that an oceanic teleconnection between AMOC strength and subsurface temperature in the EEA impacted the intensity of the WAM on millennial time scales.

Impact of abrupt deglacial climate change on tropical Atlantic subsurface temperatures.

Both instrumental data analyses and coupled ocean-atmosphere models indicate that Atlantic meridional overturning circulation (AMOC) variability is tightly linked to abrupt tropical North Atlantic (TNA) climate change through both atmospheric and oceanic processes. Although a slowdown of AMOC results in an atmospheric-induced surface cooling in the entire TNA, the subsurface experiences an even larger warming because of rapid reorganizations of ocean circulation patterns at intermediate water depths. Here, we reconstruct high-resolution temperature records using oxygen isotope values and Mg/Ca ratios in both surface- and subthermocline-dwelling planktonic foraminifera from a sediment core located in the TNA over the last 22 ky. Our results show significant changes in the vertical thermal gradient of the upper water column, with the warmest subsurface temperatures of the last deglacial transition corresponding to the onset of the Younger Dryas. Furthermore, we present new analyses of a climate model simulation forced with freshwater discharge into the North Atlantic under Last Glacial Maximum forcings and boundary conditions that reveal a maximum subsurface warming in the vicinity of the core site and a vertical thermal gradient change at the onset of AMOC weakening, consistent with the reconstructed record. Together, our proxy reconstructions and modeling results provide convincing evidence for a subsurface oceanic teleconnection linking high-latitude North Atlantic climate to the tropical Atlantic during periods of reduced AMOC across the last deglacial transition.

Age-related differences in corticomotor excitability and inhibitory processes during a visuomotor RT task.

This study tested the postulation that change in the ability to modulate corticospinal excitability and inhibitory processes underlie age-related differences in response preparation and generation during tasks requiring either rapid execution of a motor action or actively withholding that same action. Younger (n = 13, mean age = 26.0 years) and older adults (n = 13, mean age = 65.5 years) performed an RT task in which a warning signal (WS) was followed by an imperative signal (IS) to which participants were required to respond with a rapid flexion of the right thumb (go condition) or withhold their response (no-go condition). We explored the neural correlates of response preparation, generation, and inhibition using single- and paired-pulse TMS, which was administered at various times between WS and IS (response preparation phase) and between IS and onset of response-related muscle activity in the right thumb (response generation phase). Both groups exhibited increases in motor-evoked potential amplitudes (relative to WS onset) during response generation; however, this increase began earlier and was more pronounced for the younger adults in the go condition. Moreover, younger adults showed a general decrease in short-interval intracortical inhibition during response preparation in both the go and no-go conditions, which was not observed in older adults. Importantly, correlation analysis suggested that for older adults the task-related increases of corticospinal excitability and intracortical inhibition were associated with faster RT. We propose that the declined ability to functionally modulate corticospinal activity with advancing age may underlie response slowing in older adults.

Age-related differences in corticospinal excitability and inhibition during coordination of upper and lower limbs.

The ability to coordinate upper and lower limbs--a prerequisite for many everyday activities--is known to decline with age. Here we report 2 experiments in which transcranial magnetic stimulation (TMS) was used to assess corticospinal excitatory and inhibitory processes in younger and older adults during cyclical hand-foot movements. In experiment 1, motor evoked potentials (MEP) and silent period (SP) durations were measured from the active right extensor carpi radialis (ECR) muscle while it executed rhythmic oscillations in conjunction with the right or left foot. Younger adults exhibited increased SP with ipsilateral limb combinations and decreased SP with contralateral limb combinations, relative to a baseline hand only condition. Strikingly, older adults exhibited a reduced SP when ipsilateral limbs moved in opposite directions. This effect was found to be most pronounced in those older adults who exhibited poor coordination performance, suggesting that the inability to regulate inhibitory processes may underlie age-related degradation of task performance. Experiment 2 examined motor evoked potentials and SP duration in the left extensor carpi radialis which maintained a tonic contraction while the coordination task was undertaken by the right arm and right or left foot. For younger adults, coordination of ipsilateral limbs was accompanied by increased inhibition in the ipsilateral motor cortex than during the coordination of contralateral limbs. No differences in SP between conditions were noted for the older adults. In summary, older adults' reduced ability to coordinate upper and lower limbs may be related to the capacity to regulate inhibitory function in both hemispheres. This study suggests for the first time a direct link between age-related differences in interlimb coordination and the control of corticospinal inhibitory processes.

Long-lasting contralateral motor cortex excitability is increased by unilateral hand movement that triggers electrical stimulation of opposite homologous muscles.

BACKGROUND/OBJECTIVE: Long-term changes in the motor cortex can be induced by practicing motor tasks with simultaneous application of peripheral nerve stimulation. This combination may augment motor rehabilitation after stroke but has been used primarily during contraction of the affected hand. The authors tested the effect of a right hand movement that electrically stimulated left hand contraction on right motor cortex excitability. METHODS: Three tasks were used in 15 healthy subjects--a motor and stimulation task (MS(task)), stimulation only task (S(task)), and motor only task (M(task)). The MS(task) consisted of isometric thumb abduction of the right hand that triggered paired electrical stimulation of the left abductor pollicis brevis (APB) and first dorsal interosseus (FDI) motor points. The S(task) was performed 1 week later and matched the stimulation received in the MS(task). The M(task) was performed as a control. Transcranial magnetic stimulation applied to the right motor cortex assessed corticospinal excitability, short latency intracortical inhibition, and intracortical facilitation of the FDI and APB before, immediately after, and 30 minutes after task performance. RESULTS: Corticospinal excitability increased in the FDI and APB following the MS(task) but not following the S(task) or M(task). The increased excitability present 30 minutes after the MS(task) also correlated with excitability measures recorded 1 week later. CONCLUSION: A bilateral motor and electrical stimulation task can drive persistent adaptation within the corticospinal system. Hemiplegic subjects who have poor voluntary movement of the affected hand may be able to contract the unaffected hand to activate and train homologous movements.

Absence of cross-limb transfer of performance gains following ballistic motor practice in older adults.

The phenomenon of cross-limb transfer, in which unilateral strength training can result in bilateral strength gains, has recently been tested for ballistic movements. Performance gains associated with repetitive motor practice, and the associated transfer, occur within a few minutes. In this study, young and older adults were trained to perform ballistic abductions of their dominant (right) index finger as quickly as possible. Performance was assessed bilaterally before, during, and after this training. Both groups exhibited large performance gains in the right hand as a result of training (P < 0.001; young 84% improvement, older 70% improvement), which were not significantly different between groups (P = 0.40). Transcranial magnetic stimulation revealed that the performance improvements were accompanied by increases in excitability, together with decreases in intracortical inhibition, of the projections to both the trained muscle and the homologous muscle in the contralateral limb (P < 0.05). The young group also exhibited performance improvements as a result of cross-limb transfer in the left (untrained) hand (P < 0.005), equivalent to 75% of the performance increase in the trained hand. In contrast, there were no significant performance gains in the left hand for the older group (P = 0.23). This was surprising given that the older group exhibited a significantly greater degree of mirror activity than the young group (P < 0.01) in the left first dorsal interosseus muscle (FDI) during right hand movements. Our findings suggest that older adults exhibit a reduced capacity for cross-limb transfer, which may have implications for motor rehabilitation programs after stroke.

Unilateral contractions modulate interhemispheric inhibition most strongly and most adaptively in the homologous muscle of the contralateral limb.

We investigated how volitional contractions affect interhemispheric inhibition (IHI) from the active to the passive hemisphere. Younger and older adults isometrically contracted their dominant thumb (abductor pollicis brevis, APB) to various force targets. In ballistic contraction trials, transcranial magnetic stimulation (TMS) was administered very shortly after the onset of APB activity. In tonic contraction trials, TMS was delivered while the target force was maintained. In control trials both thumbs remained quiescent. In all trials, a test stimulus (TS) was directed to the APB hotspot in the non-dominant hemisphere (130% left APB resting motor threshold, RMT). In half the trials, a conditioning stimulus (130% right APB RMT) was applied to the APB hotspot in the dominant hemisphere 10 ms prior to the TS. Targeted ballistic contractions of the right APB were found to modulate (increase) IHI measured in the left APB, as previously reported for tonic contractions. Furthermore, the extent of the IHI increase was found to scale with the strength of the contralateral ballistic or tonic contraction. Less pronounced, but statistically significant, IHI increases were also observed in the left abductor digiti minimi and extensor carpi radialis during right APB contraction. For these muscles, however, the extent of the IHI modulation was independent of APB contraction strength. The capacity to modulate inhibition during contractions was unaffected by advancing age. During volitional actions, the ability to modulate IHI most adaptively in the homologous muscle of the resting limb may contribute to the prevention of mirror movements.

The effect of ballistic thumb contractions on the excitability of the ipsilateral motor cortex.

We investigated how ballistic contractions of the left thumb affect the excitability of the ipsilateral motor cortex using transcranial magnetic stimulation (TMS). TMS was applied at the motor hotspot for the right abductor pollicis brevis (APB) muscle. In 'self-triggered' trials, participants made targeted, isometric, contractions of the left APB. The right APB was either relaxed or maintained a tonic contraction. TMS was administered as soon as possible after electromyographic onset in the left APB. In 'control' trials, the left thumb remained quiescent and TMS was triggered by the computer. In each condition, 20-24 trials were conducted. Half these trials involved a single test stimulus, TS (130% APB resting motor threshold, RMT). In the other trials, short-interval intracortical inhibition (SICI) was investigated by applying a conditioning stimulus (70% APB RMT) 3 ms prior to the TS. SICI ratios were not significantly different in self-triggered and control trials. However, when the right APB was active, significantly shorter silent periods (SPs) were observed in self-triggered trials when compared with control trials. Our results support the view that SICI and SP are mediated by different inhibitory circuits, and that ipsilateral GABA(B)-ergic circuits (assessed by SP), but not GABA(A)-ergic circuits (assessed by SICI), are affected in the period immediately following voluntary ballistic contractions.

Rapid subtropical North Atlantic salinity oscillations across Dansgaard-Oeschger cycles.

Geochemical and sedimentological evidence suggest that the rapid climate warming oscillations of the last ice age, the Dansgaard-Oeschger cycles, were coupled to fluctuations in North Atlantic meridional overturning circulation through its regulation of poleward heat flux. The balance between cold meltwater from the north and warm, salty subtropical gyre waters from the south influenced the strength and location of North Atlantic overturning circulation during this period of highly variable climate. Here we investigate how rapid reorganizations of the ocean-atmosphere system across these cycles are linked to salinity changes in the subtropical North Atlantic gyre. We combine Mg/Ca palaeothermometry and oxygen isotope ratio measurements on planktonic foraminifera across four Dansgaard-Oeschger cycles (spanning 45.9-59.2 kyr ago) to generate a seawater salinity proxy record from a subtropical gyre deep-sea sediment core. We show that North Atlantic gyre surface salinities oscillated rapidly between saltier stadial conditions and fresher interstadials, covarying with inferred shifts in the Tropical Atlantic hydrologic cycle and North Atlantic overturning circulation. These salinity oscillations suggest a reduction in precipitation into the North Atlantic and/or reduced export of deep salty thermohaline waters during stadials. We hypothesize that increased stadial salinities preconditioned the North Atlantic Ocean for a rapid return to deep overturning circulation and high-latitude warming by contributing to increased North Atlantic surface-water density on interstadial transitions.

Links between salinity variation in the Caribbean and North Atlantic thermohaline circulation.

Variations in the strength of the North Atlantic Ocean thermohaline circulation have been linked to rapid climate changes during the last glacial cycle through oscillations in North Atlantic Deep Water formation and northward oceanic heat flux. The strength of the thermohaline circulation depends on the supply of warm, salty water to the North Atlantic, which, after losing heat to the atmosphere, produces the dense water masses that sink to great depths and circulate back south. Here we analyse two Caribbean Sea sediment cores, combining Mg/Ca palaeothermometry with measurements of oxygen isotopes in foraminiferal calcite in order to reconstruct tropical Atlantic surface salinity during the last glacial cycle. We find that Caribbean salinity oscillated between saltier conditions during the cold oxygen isotope stages 2, 4 and 6, and lower salinities during the warm stages 3 and 5, covarying with the strength of North Atlantic Deep Water formation. At the initiation of the Bølling/Allerød warm interval, Caribbean surface salinity decreased abruptly, suggesting that the advection of salty tropical waters into the North Atlantic amplified thermohaline circulation and contributed to high-latitude warming.

Direction of foot force for pushes against a fixed pedal: role of effort level.

Control of the force exerted by the foot on the ground is critical to human locomotion. During running on a treadmill and pushing against a fixed pedal, humans increased foot force in a linear manner in sagittal plane force space. However, for pushes against a moving pedal, force output was linear for some participants but slightly curved for others. A primary difference between the static and dynamic pedaling studies was that the dynamic study required participants to push with varying peak effort levels, whereas a constant peak effort level was used for the fixed pedal pushes. The present study evaluated the possibility that force direction varied with level of effort. Seated humans pushed against a fixed pedal to a series of force magnitude targets. The force direction varied systematically with effort level consistent with the force path curvature observed for dynamic pedaling.

The control of foot force during pushing efforts against a moving pedal.

The muscle component of the force applied to a bicycle pedal (foot force) by seated humans provided insight into the organization of the motor system. Healthy adults ( n=11) pedaled a stationary cycle ergometer while attempting to match peak foot force magnitude to visually presented force targets (200, 250,., 650 N). Pedaling cadence was maintained at 60 rpm by a motor. Measurements of the foot force, pedal angle, and crank angle were recorded. The experimental design and data analysis allowed the isolation of the muscle component of the foot force from the contributions due to gravity and inertia. A graphical representation of the muscle component of the foot force (force path) was created for each of several crank angles throughout the extension phase of the pedaling cycle. The force paths showed several highly conserved characteristics across participants and crank angles. Each force path occupied a narrow range in force space despite the ability of the participants to produce force in a wide region of force space. Three control strategies were observed in the geometry of the force paths. Eighty five percent of the force paths were linear for six of the participants, and 79% of the force paths had second-order curvature for the other five participants. The curvature was concave to the posterior for four of the participants and concave to the anterior for one participant. The linear force paths were consistent with the previously reported linear nature of the force paths for pushes against a quasi-static pedal. The observation of simple force path geometry for two tasks with dissimilar dynamic characteristics suggests that this aspect of foot force control may be common to a range of lower limb tasks and may reflect a mechanism by which the nervous system organizes the control of foot force.

Direction of foot force for pushes against a fixed pedal: variation with pedal position.

The force that healthy humans generated against a fixed pedal was measured and compared with that predicted by four models. The participants (n = 11) were seated on a stationary bicycle and performed brief pushing efforts against an instrumented pedal with the crank fixed. Pushes were performed to 10 force magnitude targets and at 12 crank angles. The increasing magnitude portion of the sagittal-plane force path for each push effort was fitted with a line to determine the direction of the muscle component of the foot force. Those directions varied systematically with the position of the pedal (crank angle) such that the force path lines intersected a common region superior and slightly anterior to the hip. The ability of four models to predict force path direction was tested. All four models captured the general variation of direction with pedal position. Two of the models provided the best performance. One was a musculoskeletal model consisting of nine muscles with parameters adjusted to provide the best possible fit. The other model was derived from (a) observations that the lines-of-action of the muscle component of foot force tended to intersect in a common region near the hip, and (b) the corresponding need for foot force to intersect the center-of-mass during walking. Thus, this model predicted force direction at each pedal position as that of a line intersecting the pedal pivot and a common point located near the hip (divergent point). The results suggest that the control strategy employed in this seated pushing task reflects the extensive experience of the leg in directing force appropriately to maintain upright posture and that relative muscle strengths have adapted to that pattern of typical activation.