Noise analysis and relaxation experiments of transport of hydrophobic anions across lipid membranes at equilibrium and nonequilibrium

Under equilibrium and nonequilibnum steady-stale conditions the spectral intensity of current noise S_J(f) generated by the transport of hydrophobic unions across lipid bilayer membranes was investigated. The experimental results were compared with different reaction models S_J(f) showed a characteristic increase proportional to f² between frequency-independent tails at low and high frequencies. This gradient was found to be independent of applied voltage which indicates the contribution of a single voltage-dependent reaction step of ion translocation across the membrane From the shape of S_J(f) at low frequencies the rate constant of ion desorption from the membrane into the aqueous phase could be estimated. Unambiguous evidence for the application of a general model, which includes the coupling of slow ion diffusion in the aqueous phase to ion adsorption/desorption at the membrane interface, could not be obtained from the low-frequency shape of S_J(f). The shot noise of this ion transport determines the amplitude of S_J(f) at high frequencies which decreases with increasing voltage applied. Analysis of voltage-jump current-relaxation experiments and of current noise carried cut on one membrane yielded significant differences of the derived ion partition coefficient. This deviation is qualitatively described on the basis of incomplete reaction steps.     

Activation of Shaker Potassium Channels : I. Characterization of Voltage-dependent Transitions

The conformational changes associated with activation gating in Shaker potassium channels are functionally characterized in patch-clamp recordings made from Xenopus laevis oocytes expressing Shaker channels with fast inactivation removed. Estimates of the forward and backward rates for transitions are obtained by fitting exponentials to macroscopic ionic and gating current relaxations at voltage extremes, where we assume that transitions are unidirectional. The assignment of different rates is facilitated by using voltage protocols that incorporate prepulses to preload channels into different distributions of states, yielding test currents that reflect different subsets of transitions. These data yield direct estimates of the rate constants and partial charges associated with three forward and three backward transitions, as well as estimates of the partial charges associated with other transitions. The partial charges correspond to an average charge movement of 0.5 e₀ during each transition in the activation process. This value implies that activation gating involves a large number of transitions to account for the total gating charge displacement of 13 e₀. The characterization of the gating transitions here forms the basis for constraining a detailed gating model to be described in a subsequent paper of this series.     

Dynamic characterization of human breast cancer cells using a piezoresistive microcantilever

In this paper, frequency response (dynamic compression and recovery) is suggested as a new physical marker to differentiate between breast cancer cells (MCF7) and normal cells (MCF10A). A single cell is placed on the laminated piezoelectric actuator and a piezoresistive microcantilever is placed on the upper surface of the cell at a specified preload displacement (or an equivalent force). The piezoelectric actuator excites the single cell in a sinusoidal fashion and its dynamic deformation is then evaluated from the displacement converted by measuring the voltage output through a piezoresistor in the microcantilever. The microcantilever has a flat contact surface with no sharp tip, making it possible to measure the overall properties of the cell rather than the local properties. These results indicate that the MCF7 cells are more deformable in quasi-static conditions compared with MCF10A cells, consistent with known characteristics. Under conditions of high frequency of over 50 Hz at a 1 μm preload displacement, 1 Hz at a 2 μm preload displacement, and all frequency ranges tested at a 3 μm preload displacement, MCF7 cells showed smaller deformation than MCF10A cells. MCF7 cells have higher absorption than MCF10A cells such that MCF7 cells appear to have higher deformability according to increasing frequency. Moreover, larger preload and higher frequencies are shown to enhance the differences in cell deformability between the MCF7 cells and MCF10A cells, which can be used as a physical marker for differentiating between MCF10A cells and MCF7 cells, even for high-speed screening devices.     

Biofilm thickness measurements by light microscopy

Light microscopy has been used to measure biofilm thickness. The vertical displacement of the sample required to focus from the biofilm-liquid interface to the biofilm-substratum interface is measured by the stage micrometer. Biofilm thickness is proportional, but not equal, to the measured vertical displacement. An expression for the proportionality constant, k_f, in terms of refractive indices is determined from a geometric analysis of the light path. k_f can be estimated as the ratio of the refractive index of the film to the refractive index of the media interfacing the film between the objective lens and the sample. The thickness of any transparent film may be determined by light microscopy when the refractive index of the film is known.     

Modeling combined transport of water and charged graded-size molecules across the glomerular capillary wall

Clearance studies using various probe molecules established that the passage of molecules/proteins across the glomerular capillary wall of mammalian kidneys is increasingly restricted as their size and net negative charge increase. An extended mathematical model, based on the Fiber Matrix theory, was developed to describe the dynamics of the size- and charge-selective functions of the glomerular capillary barrier using mainly its hemodynamic, morphometric, and electrostatic variables. The glomerular basement membrane was represented as a homogeneous three-dimensional network of fibers of uniform length (L_f), radius (R_f), and packing density (N_fv) and characteristic Darcy permeability. The model was appropriate for simulating fractional clearance data of neutral and charged solutes from an experimental modeling exercise. We believe that the L_f and R_f best-fit numerical values may signify new insights for the diagnosis of some human nephropathies.     

A computational study of invariant I5 in a nearly incompressible transversely isotropic model for white matter

The aligned axonal fiber bundles in white matter make it suitable to be modeled as a transversely isotropic material. Recent experimental studies have shown that a minimal form, nearly incompressible transversely isotropic (MITI) material model, is capable of describing mechanical anisotropy of white matter. Here, we used a finite element (FE) computational approach to demonstrate the significance of the fifth invariant (I₅) when modeling the anisotropic behavior of white matter in the large-strain regime. We first implemented and validated the MITI model in an FE simulation framework for large deformations. Next, we applied the model to a plate-hole structural problem to highlight the significance of the invariant I₅ by comparing with the standard fiber reinforcement (SFR) model. We also compared the two models by fitting the experiment data of asymmetric indentation, shear test, and uniaxial stretch of white matter. Our results demonstrated the significance of I₅ in describing shear deformation/anisotropy, and illustrated the potential of the MITI model to characterize transversely isotropic white matter tissues in the large-strain regime.     

Theory and Simulation of Water Permeation in Aquaporin-1

We discuss the difference between osmotic permeability p_f and diffusion permeability p_d of single-file water channels and demonstrate that the p_f/p_d ratio corresponds to the number of effective steps a water molecule needs to take to permeate a channel. While p_d can be directly obtained from equilibrium molecular dynamics simulations, p_f can be best determined from simulations in which a chemical potential difference of water has been established on the two sides of the channel. In light of this, we suggest a method to induce in molecular dynamics simulations a hydrostatic pressure difference across the membrane, from which p_f can be measured. Simulations using this method are performed on aquaporin-1 channels in a lipid bilayer, resulting in a calculated p_f of 7.1 × 10⁻¹⁴ cm³/s, which is in close agreement with observation. Using a previously determined p_d value, we conclude that p_f/p_d for aquaporin-1 measures ~12. This number is explained in terms of channel architecture and conduction mechanism.     

Cytochrome b6f – Orchestrator of photosynthetic electron transfer

Cytochrome b₆f (cytb₆f) lies at the heart of the light-dependent reactions of oxygenic photosynthesis, where it serves as a link between photosystem II (PSII) and photosystem I (PSI) through the oxidation and reduction of the electron carriers plastoquinol (PQH₂) and plastocyanin (Pc). A mechanism of electron bifurcation, known as the Q-cycle, couples electron transfer to the generation of a transmembrane proton gradient for ATP synthesis. Cytb₆f catalyses the rate-limiting step in linear electron transfer (LET), is pivotal for cyclic electron transfer (CET) and plays a key role as a redox-sensing hub involved in the regulation of light-harvesting, electron transfer and photosynthetic gene expression. Together, these characteristics make cytb₆f a judicious target for genetic manipulation to enhance photosynthetic yield, a strategy which already shows promise. In this review we will outline the structure and function of cytb₆f with a particular focus on new insights provided by the recent high-resolution map of the complex from Spinach.     

Effect of deuterium oxide on contraction characteristics and ATPase activity in glycerinated single rabbit skeletal muscle fibers

We studied the effect of deuterium oxide (D₂O) on contraction characteristics and ATPase activity of single glycerinated muscle fibers of rabbit psoas. D₂O increased the maximum isometric force P₀ by about 20%, while the force versus stiffness relation did not change appreciably. The maximum shortening velocity under zero load V_max did not change appreciably in D₂O, so that the force-velocity (P-V) curve was scaled depending on the value of P₀. The Mg-ATPase activity of the fibers during generation of steady isometric force P₀ was reduced by about 50% in D₂O. Based on the Huxley contraction model, these results can be accounted for in terms of D₂O-induced changes in the rate constants f₁ and g₁ for making and breaking actin-myosin linkages in the isometric condition, in such a way that f₁/(f₁+g₁) increases by about 20%, while (f₁+g₁) remains unchanged. The D₂O effect at the molecular level is discussed in connection with biochemical studies on actomyosin ATPase.     

X-ray dose reduction using additional copper filtration for dental cone beam CT

This study aims to quantitatively evaluate the effect of additional copper-filters (Cu-filters) on the radiation dose and contrast-to-noise ratio (CNR) in a dental cone beam computed tomography (CBCT). The Cu-filter thickness and tube voltage of the CBCT unit were varied in the range of 0.00–0.20 mm and 70–90 kV, respectively. The CBCT images of a phantom with homogeneous materials of aluminum, air, and bone equivalent material (BEM) were acquired. The CNRs were calculated from the voxel values of each homogeneous material. The CTDI_vol was measured using standard polymethyl methacrylate CTDI test objects. We evaluated and analyzed the effects of tube current and various radiation qualities on the CNRs and CTDI_vol. We observed a tendency for higher CNR at increasing tube voltage and tube current in all the homogeneous materials. On the other hand, the CNR reduced at increasing Cu-filter thickness. The tube voltage of 90 kV showed a clear advantage in the tube current–CNR curves in all the homogeneous materials. The CTDI_vol increased as the tube voltage and tube current increased and decreased with the increase in the Cu-filter thickness. When the CNR was fixed at 9.23 of BEM at an exposure setting of 90 kV/5 mA without a Cu-filter, the CTDI_vol at 90 kV with Cu-filters was 8.7% lower compared with that at 90 kV without a Cu-filter. The results from this study demonstrate the potential of adding a Cu-filter for patient dose reduction while ensuring the image quality.     

Plasmon-Matter Interactions in Optoelectronic Metamaterials with Negative Refractive Index

Optoelectronic metamaterials composed of nanoscale metallic structures and semiconductor quantum structures constitute a powerful platform to explore light-matter interaction and new devices. In this work, we numerically study an optoelectronically coupled metamaterial consisting of metallic double fishnet (DF) layers and semiconductor quantum well (QW) spacing layer. When the electronic intersubband transition (ISBT) in the QW coincides with the plasmonic resonances of the DF structure, the plasmon-matter interaction (PMI) can modify the optical properties considerably. In case of the ISBT-matching localized surface plasmons (LSP), i.e., f _QW?=?f _LSP, the polarization-selection-rule forbidden ISBT absorption can be enabled due to the nonnegligible E _z ?field distributions while the retrieved optical constants remain almost unchanged. However, when the gap surface plasmons (GSP) are matched, i.e., f _QW?=?f _GSP, PMI exhibits a clear anti-crossing behavior implying strong coupling effects between ISBT and GSP resonance and formation of intersubband polaritons. The effective optical constants are therefore modulated appreciably. The large difference between GSP and LSP can be attributed to their distinctive resonance qualities (Q-factors) and polarization conversion ratios (99.28 % for GSP and 1.54 % for LSP) from the transverse electric (TE) to transverse magnetic (TM) mode. Our results provide insight into the physical mechanism of PMI in nanoscale semiconductor-plasmon hybrid systems and suggest an alternative means in tunable negative refractive index (NRI) applications.     

C-Terminal Modulatory Domain Controls Coupling of Voltage-Sensing to?Pore Opening in Cav1.3 L-type Ca2+ Channels

Activity of voltage-gated Ca_v1.3 L-type Ca²⁺ channels is required for proper hearing as well as sinoatrial node and brain function. This critically depends on their negative activation voltage range, which is further fine-tuned by alternative splicing. Shorter variants miss a C-terminal regulatory domain (CTM), which allows them to activate at even more negative potentials than C-terminally long-splice variants. It is at present unclear whether this is due to an increased voltage sensitivity of the Ca_v1.3 voltage-sensing domain, or an enhanced coupling of voltage-sensor conformational changes to the subsequent opening of the activation gate. We studied the voltage-dependence of voltage-sensor charge movement (Q_ON-V) and of current activation (I_Ca-V) of the long (Ca_v1.3_L) and a short Ca_v1.3 splice variant (Ca_v1.3_42A) expressed in tsA-201 cells using whole cell patch-clamp. Charge movement (Q_ON) of Ca_v1.3_L displayed a much steeper voltage-dependence and a more negative half-maximal activation voltage than Ca_v1.2 and Ca_v3.1. However, a significantly higher fraction of the total charge had to move for activation of Ca_v1.3 half-maximal conductance (Ca_v1.3: 68%; Ca_v1.2: 52%; Ca_v3.1: 22%). This indicated a weaker coupling of Ca_v1.3 voltage-sensor charge movement to pore opening. However, the coupling efficiency was strengthened in the absence of the CTM in Ca_v1.3_42A, thereby shifting I_Ca-V by 7.2 mV to potentials that were more negative without changing Q_ON-V. We independently show that the presence of intracellular organic cations (such as n-methyl-D-glucamine) induces a pronounced negative shift of Q_ON-V and a more negative activation of I_Ca-V of all three channels. These findings illustrate that the voltage sensors of Ca_v1.3 channels respond more sensitively to depolarization than those of Ca_v1.2 or Ca_v3.1. Weak coupling of voltage sensing to pore opening is enhanced in the absence of the CTM, allowing short Ca_v1.3_42A splice variants to activate at lower voltages without affecting Q_ON-V.     

Sperm competition games: Sperm size (mass) and number under raffle and displacement, and the evolution of P2

We examine models for evolution of sperm size (i.e. mass m) and number (s) under three mechanisms of sperm competition at low ‘risk’ levels: (i) raffle with no constraint on space available for competing sperm, (ii) direct displacement mainly by seminal fluid, and (iii) direct displacement mainly by sperm mass. Increasing sperm mass increases a sperm's ‘competitive weight’ against rival sperm through a diminishing returns function, r(m). ESS total ejaculate expenditure (the product m^?s^?) increases in all three models with sperm competition risk, q. If r(m), or ratio r′(m)/r(m), is independent of ESS sperm numbers, ESS sperm mass remains constant, and the sperm mass/number ratio (m^?/s^?) therefore decreases with risk. Dependency of sperm mass on risk can arise if r(m) depends on competing sperm density (sperm number / space available for sperm competition). Such dependencies generate complex relationships between sperm mass and number with risk, depending both on the mechanism and how sperm density affects r(m). While numbers always increase with risk, mass can either increase or decrease, but m^?/s^? typically decreases with risk unless sperm density strongly influences r(m). Where there is no extrinsic loading due to mating order, ESS paternity of the second (i.e. last) male to mate (P₂) under displacement always exceeds 0.5, and increases with risk (in the raffle P₂=0.5). Caution is needed when seeking evidence for a sperm size-number trade off. Although size and number trade-off independently against effort spent on acquiring matings, their product, m^?s^?, is invariant or fixed at a given risk level, effectively generating a size-number trade off. However, unless controlled for the effects of risk, the relation between m^? and s^? can be either positive or negative (a positive relation is usually taken as evidence against a size-number trade off).     

C-Terminal Modulatory Domain Controls Coupling of Voltage-Sensing to Pore Opening in Cav1.3 L-type Ca Channels

Activity of voltage-gated Ca_v1.3 L-type Ca²⁺ channels is required for proper hearing as well as sinoatrial node and brain function. This critically depends on their negative activation voltage range, which is further fine-tuned by alternative splicing. Shorter variants miss a C-terminal regulatory domain (CTM), which allows them to activate at even more negative potentials than C-terminally long-splice variants. It is at present unclear whether this is due to an increased voltage sensitivity of the Ca_v1.3 voltage-sensing domain, or an enhanced coupling of voltage-sensor conformational changes to the subsequent opening of the activation gate. We studied the voltage-dependence of voltage-sensor charge movement (Q_ON-V) and of current activation (I_Ca-V) of the long (Ca_v1.3_L) and a short Ca_v1.3 splice variant (Ca_v1.3_42A) expressed in tsA-201 cells using whole cell patch-clamp. Charge movement (Q_ON) of Ca_v1.3_L displayed a much steeper voltage-dependence and a more negative half-maximal activation voltage than Ca_v1.2 and Ca_v3.1. However, a significantly higher fraction of the total charge had to move for activation of Ca_v1.3 half-maximal conductance (Ca_v1.3: 68%; Ca_v1.2: 52%; Ca_v3.1: 22%). This indicated a weaker coupling of Ca_v1.3 voltage-sensor charge movement to pore opening. However, the coupling efficiency was strengthened in the absence of the CTM in Ca_v1.3_42A, thereby shifting I_Ca-V by 7.2 mV to potentials that were more negative without changing Q_ON-V. We independently show that the presence of intracellular organic cations (such as n-methyl-D-glucamine) induces a pronounced negative shift of Q_ON-V and a more negative activation of I_Ca-V of all three channels. These findings illustrate that the voltage sensors of Ca_v1.3 channels respond more sensitively to depolarization than those of Ca_v1.2 or Ca_v3.1. Weak coupling of voltage sensing to pore opening is enhanced in the absence of the CTM, allowing short Ca_v1.3_42A splice variants to activate at lower voltages without affecting Q_ON-V.     

Thermodynamic and kinetic properties of amino-terminal and S4-S5 loop HERG channel mutants under steady-state conditions

Gating kinetics and underlying thermodynamic properties of human ether-a-go-go-related gene (HERG) K⁺ channels expressed in Xenopus oocytes were studied using protocols able to yield true steady-state kinetic parameters. Channel mutants lacking the initial 16 residues of the amino terminus before the conserved eag/PAS region showed significant positive shifts in activation voltage dependence associated with a reduction of z_g values and a less negative ΔG_o, indicating a deletion-induced displacement of the equilibrium toward the closed state. Conversely, a negative shift and an increased ΔG_o, indicative of closed-state destabilization, were observed in channels lacking the amino-terminal proximal domain. Furthermore, accelerated activation and deactivation kinetics were observed in these constructs when differences in driving force were considered, suggesting that the presence of distal and proximal amino-terminal segments contributes in wild-type channels to specific chemical interactions that raise the energy barrier for activation. Steady-state characteristics of some single point mutants in the intracellular loop linking S4 and S5 helices revealed a striking parallelism between the effects of these mutations and those of the amino-terminal modifications. Our data indicate that in addition to the recognized influence of the initial amino-terminus region on HERG deactivation, this cytoplasmic region also affects activation behavior. The data also suggest that not only a slow movement of the voltage sensor itself but also delaying its functional coupling to the activation gate by some cytoplasmic structures possibly acting on the S4-S5 loop may contribute to the atypically slow gating of HERG.     

OroSTIFF: Face-referenced measurement of perioral stiffness in health and disease

A new device and automated measurement technology known as OroSTIFF is described to characterize non-participatory perioral stiffness in healthy adults for eventual application to patients with orofacial movement disorders associated with neuromotor disease, traumatic injury, or congenital clefts of the upper lip. Previous studies of perioral biomechanics required head stabilization for extended periods of time during measurement, which precluded sampling patients with involuntary body/head movements (dyskinesias), or pediatric subjects. The OroSTIFF device is face-referenced and avoids the complications associated with head-restraint. Supporting data of non-participatory perioral tissue stiffness using OroSTIFF are included from 10 male and 10 female healthy subjects. The OroSTIFF device incorporates a pneumatic glass air cylinder actuator instrumented for pressure, and an integrated subminiature displacement sensor to encode lip aperture. Perioral electromyograms were simultaneously sampled to confirm passive muscle state for the superior and inferior divisions of the orbicularis oris muscles. Perioral stiffness, derived as a quotient from resultant force (ΔF) and interangle span (ΔX), was modeled with multilevel regression techniques. Real-time calculation of the perioral stiffness function demonstrated a significant quadratic relation between imposed interangle stretch and resultant force. This stiffness growth function also differed significantly between males and females. This study demonstrates the OroSTIFF ‘proof-of-concept’ for cost-effective non-invasive stimulus generation and derivation of perioral stiffness in a group of healthy unrestrained adults, and a case study to illustrate the dose-dependent effects of Levodopa on perioral stiffness in an individual with advanced Parkinson’s disease who exhibited marked dyskinesia and rigidity.     

Cervical spine posteroanterior stiffness differs with neck position

PurposeSpinal stiffness is commonly considered when treating patients with neck pain, but there are few studies reporting the objective measurement of cervical spine stiffness or the possible kinesiological factors that may affect its quantification. The aim of this study was to determine if the position of the neck affects cervical spine stiffness.MethodsAn instrumented stiffness assessment device measured posteroanterior cervical spine stiffness at C4 of 25 prone-lying asymptomatic subjects in three neck positions in randomised order: maximal flexion, maximal extension, and neutral. The device applied five standardised mechanical oscillatory pressures while measuring the applied force and concurrent displacement, defining stiffness as the slope of the linear portion of the force–displacement curve. Repeated measures analysis of variance with Bonferroni-adjusted post hoc comparisons determined whether stiffness differed between neck positions.ResultsThere was a significant difference in cervical spine stiffness between different neck positions (F_(1.6,38.0) = 16.6, P < 0.001). Stiffness was least in extension with a mean of 3.09 N/mm (95% CI 2.59, 3.58) followed by neutral (3.94, 95% CI 3.49, 4.39), and then flexion (4.32, 95% CI 3.96, 4.69).ConclusionWhen assessing cervical spine stiffness, neck position should be standardised to ensure maximal reliability and utility of stiffness judgments.     

Effects of Epinephrine on the Pacemaker Potassium Current of Cardiac Purkinje Fibers

Epinephrine promotes spontaneous activity in cardiac Purkinje fibers through its action on the pacemaker potassium current (i_KK2). The mechanism of the acceleratory effect was studied by means of a voltage clamp technique. The results showed that the hormone speeds the deactivation of i_KK2 during pacemaker activity by displacing the kinetic parameters of i_KK2 toward less negative potentials. This depolarizing voltage shift is the sole explanation of the acceleratory effect since epinephrine did not alter the rectifier properties of i_KK2, or the underlying inward leakage current, or the threshold for i_NNa. The dose dependence of the voltage shift in the i_KK2 activation curve was similar in 1.8 and 5.4 mM [Ca]_o. The maximal voltage shift (usually ~20 mV) was produced by epinephrine concentrations of > 10^-6 M. The half-maximal effect was evoked by 60 nM epinephrine, nearly an order of magnitude lower than required for half-maximal effect on the secondary inward current (Carmeliet and Vereecke, 1969). The β-blocker propranolol (10^-6 M) prevented the effect of epinephrine (10^-7M) but by itself gave no voltage shift. Epinephrine shifted the activation rate coefficient α₈ to a greater extent than the deactivation rate coefficient β₈, and often steepened the voltage dependence of the steady-state activation curve. These deviations from simple voltage shift behavior were discussed in terms of possible mechanisms of epinephrine's action on the i_KK2 channel.     

Voltage clamp studies of fertilization in sea urchin eggs: I. Effect of clamped membrane potential on sperm entry,activation, and development

To analyze the role of the activation potential (a positive shift of the membrane potential which occurs following sperm attachment) in fertilization and development of the sea urchin egg, unfertilized Lytechinus variegatus eggs were voltage clamped at membrane potentials (E_m) from +20 to ?90 mV, and then inseminated. Either a fast two electrode voltage clamp, or a single electrode switched voltage clamp was used. The clamp was maintained for 3 to 15 min after initiation of a conductance increase. At E_m more positive than +18 mV, even though many sperm may attach, the egg remains completely inert (Jaffe, Nature (London)261, 68–71, 1976). At E_m from +17 to ?90 mV, all inseminated eggs elevate normal fertilization envelopes, although substantially increased concentrations of sperm are required at E_m from +17 to +12 mV. Whether cleavage occurs depends on the clamped E_m. When clamped at E_m from +17 to ?25 mV, 100% of activated eggs cleave. However, when clamped at E_m from ?26 to ?75 mV the percentage of activated eggs which cleave progressively decreases. At clamped E_m between ?76 and ?90 mV, none of the activated eggs cleave. All monospermic voltage clamped eggs that cleave develop to normal swimming blastulae. In all eggs that fail to cleave (clamped at E_m more negative than ?30 mV), sperm penetration is blocked, the sperm is lifted off the egg surface as the fertilization envelope rises, and a sperm aster never forms. Preventing formation of the fertilization envelope by prior disruption of the vitelline layer with dithiothreitol does not promote entry of the sperm. In conclusion, preventing the depolarization normally associated with fertilization suppresses sperm entry in the sea urchin egg, yet activation proceeds. Present evidence suggests an effect of the electrical field across the plasma membrane in suppressing sperm entry.     

Heliobacterial Rieske/cytb complex

Data on structure and function of the Rieske/cytb complex from Heliobacteria are scarce. They indicate that the complex is related to the b ₆ f complex in agreement with the phylogenetic position of the organism. It is composed of a diheme cytochrome c, and a Rieske iron–sulfur protein, together with transmembrane cytochrome b ₆ and subunit IV. Additional small subunits may be part of the complex. The cofactor content comprises heme c _i, first discovered in the Q_i binding pocket of b ₆ f complexes. The redox midpoint potentials are more negative than in b ₆ f complex in agreement with the lower redox midpoint potentials (by about 150 mV) of its reaction partners, menaquinone, and cytochrome c ₅₅₃. The enzyme is implicated in cyclic electron transfer around the RCI. Functional studies are favored by the absence of antennae and the simple photosynthetic reaction chain but are hampered by the high oxygen sensitivity of the organism, its chlorophyll, and lipids.