Research article Topical Sections

STIM1 activation is regulated by a 14 amino acid sequence adjacent to the CRAC activation domain

  • Received: 19 December 2015 Accepted: 24 February 2016 Published: 28 February 2016
  • Oligomerization of the Ca2+ sensor, STIM1, in the endoplasmic reticulum (ER) membrane, caused by depletion of ER Ca2+ stores, results in STIM1 coupling to the plasma membrane Ca2+ channel protein, Orai1, to activate Ca2+ influx in a process known as store-operated Ca2+ entry. We use fluorimetry-based fluorescence resonance energy transfer (FRET) to monitor changes in STIM1 oligomerization in COS7 cells transfected with STIM1 constructs containing selected truncations, deletions, and point mutations, and labeled with donor and acceptor fluorescent proteins at either the luminal (N-terminal) or the cytoplasmic (C-terminal) ends. Our results with sequential truncations of STIM1 from the C-terminus support previous evidence that the CRAC activation domain (CAD/SOAR, human sequence 342–448) is an oligomer-promoting segment of STIM1, and they show that truncation just after CAD/SOAR (1–448) causes significantly elevated basal cytoplasmic Ca2+ and spontaneous STIM1 clustering. We find that a 14 amino acid sequence just C-terminal of CAD/SOAR (449–462) prevents spontaneous clustering and activation of STIM1 in COS7 cells. In response to store depletion, C-terminally labeled STIM1 without CAD/SOAR clusters together with CAD/SOAR-containing STIM1 constructs. However, these donor-acceptor pairs do not undergo a stimulated increase in FRET, exhibiting instead a decrease in FRET consistent with a stimulated conformational extension in full length STIM1. We find that the 14 amino acid sequence plays a regulatory role in this process. Overall, our FRET results provide evidence in live cells that Ca2+ store depletion stimulates a conformational extension in the cytoplasmic segment of STIM1 that accompanies its oligomerization.

    Citation: Marek K. Korzeniowski, Barbara Baird, David Holowka. STIM1 activation is regulated by a 14 amino acid sequence adjacent to the CRAC activation domain[J]. AIMS Biophysics, 2016, 3(1): 99-118. doi: 10.3934/biophy.2016.1.99

    Related Papers:

    [1] Kushan Tharuka Lulbadda, K. T. M. U. Hemapala . The additional functions of smart inverters. AIMS Energy, 2019, 7(6): 971-988. doi: 10.3934/energy.2019.6.971
    [2] J. Appelbaum . Shading and masking affect the performance of photovoltaic systems—a review. AIMS Energy, 2019, 7(1): 77-87. doi: 10.3934/energy.2019.1.77
    [3] Hari Charan Nannam, Atanu Banerjee . A novel control technique for a single-phase grid-tied inverter to extract peak power from PV-Based home energy systems. AIMS Energy, 2021, 9(3): 414-445. doi: 10.3934/energy.2021021
    [4] Stefano Farnesi, Mario Marchesoni, Massimiliano Passalacqua, Luis Vaccaro . Soft-switching cells for Modular Multilevel Converters for efficient grid integration of renewable sources. AIMS Energy, 2019, 7(3): 246-263. doi: 10.3934/energy.2019.3.246
    [5] Abdulrahman J. Babqi, NasimUllah, Ahmed Althobaiti, Hend I. Alkhammash, Asier Ibeas . Current model predictive fault-tolerant control for grid-connected photovoltaic system. AIMS Energy, 2022, 10(2): 273-291. doi: 10.3934/energy.2022015
    [6] M. A. Kalhan S. Boralessa, H. V. Vimukkthi Priyadarshana, K. T. M. Udayanga Hemapala . A review on control systems for fast demand response for ancillary services. AIMS Energy, 2020, 8(6): 1108-1126. doi: 10.3934/energy.2020.6.1108
    [7] David Ludwig, Christian Breyer, A.A. Solomon, Robert Seguin . Evaluation of an onsite integrated hybrid PV-Wind power plant. AIMS Energy, 2020, 8(5): 988-1006. doi: 10.3934/energy.2020.5.988
    [8] Sabir Rustemli, Zeki İlcihan, Gökhan Sahin, Wilfried G. J. H. M. van Sark . A novel design and simulation of a mechanical coordinate based photovoltaic solar tracking system. AIMS Energy, 2023, 11(5): 753-773. doi: 10.3934/energy.2023037
    [9] Guy M. Toche Tchio, Joseph Kenfack, Joseph Voufo, Yves Abessolo Mindzie, Blaise Fouedjou Njoya, Sanoussi S. Ouro-Djobo . Diagnosing faults in a photovoltaic system using the Extra Trees ensemble algorithm. AIMS Energy, 2024, 12(4): 727-750. doi: 10.3934/energy.2024034
    [10] Antonino Laudani, Gabriele Maria Lozito, Valentina Lucaferri, Martina Radicioni, Francesco Riganti Fulginei . On circuital topologies and reconfiguration strategies for PV systems inpartial shading conditions: A review. AIMS Energy, 2018, 6(5): 735-763. doi: 10.3934/energy.2018.5.735
  • Oligomerization of the Ca2+ sensor, STIM1, in the endoplasmic reticulum (ER) membrane, caused by depletion of ER Ca2+ stores, results in STIM1 coupling to the plasma membrane Ca2+ channel protein, Orai1, to activate Ca2+ influx in a process known as store-operated Ca2+ entry. We use fluorimetry-based fluorescence resonance energy transfer (FRET) to monitor changes in STIM1 oligomerization in COS7 cells transfected with STIM1 constructs containing selected truncations, deletions, and point mutations, and labeled with donor and acceptor fluorescent proteins at either the luminal (N-terminal) or the cytoplasmic (C-terminal) ends. Our results with sequential truncations of STIM1 from the C-terminus support previous evidence that the CRAC activation domain (CAD/SOAR, human sequence 342–448) is an oligomer-promoting segment of STIM1, and they show that truncation just after CAD/SOAR (1–448) causes significantly elevated basal cytoplasmic Ca2+ and spontaneous STIM1 clustering. We find that a 14 amino acid sequence just C-terminal of CAD/SOAR (449–462) prevents spontaneous clustering and activation of STIM1 in COS7 cells. In response to store depletion, C-terminally labeled STIM1 without CAD/SOAR clusters together with CAD/SOAR-containing STIM1 constructs. However, these donor-acceptor pairs do not undergo a stimulated increase in FRET, exhibiting instead a decrease in FRET consistent with a stimulated conformational extension in full length STIM1. We find that the 14 amino acid sequence plays a regulatory role in this process. Overall, our FRET results provide evidence in live cells that Ca2+ store depletion stimulates a conformational extension in the cytoplasmic segment of STIM1 that accompanies its oligomerization.


    1. Introduction

    Increasing efficiencies of different cell technologies and decreasing cost of installations attracted photovoltaic (PV) systems for many applications [1]. For example, they became a mainstream power generation source bringing the total world capacity to 178 GW by 2014 [2]. Another area where PV applications are increasing is the electric car industry. In 2014, Ford has announced there concept car which has rooftop PV panels with special concentrators to charge the battery. The company said that the PV panels could provide power equal to a four-hour battery charge. Eindhoven University of Technology in Holland also announced their lightweight, wedge-shaped electric car that charges itself with PV cells. Rooftop PV panels on a car reduce the transfer of heat into the cabin by solar radiation in hot climates; thus reducing the heating of air temperature inside the car. This reduces the workload of the air conditioner, resulting in reduced fuel consumption (in a hybrid). Even though these cars provide a number of benefits, the efficiency of roof top PV panels will reduce with shading: as the shading is a dynamic process, an effective way of overcoming effects of shading is quite important.

    Commonly used motors for hybrid and fully-electric vehicles are dc brushless and induction motors. They use motors having similar stators and are connected to the dc bus of the car through an inverter, usually operating on Pulse Width Modulation (PWM). This inverter requires a high voltage capacitor at the dc link and bulky filtering arrangements to eliminate unwanted harmonics. In order to overcome the limitations of PWM inverters and to facilitate PV connection, Imtiaz et. al (2013) proposed a celllevel power conversion topology based on a cascaded multi-level inverter [3]. In this inverter, a number of PV cells are connected together through H-bridge inverters. The outputs of each of different level H-bridge inverters are connected in series such that the synthesised waveform is almost a sinusoidal. Therefore the output has less harmonic distortion and switching losses are less compared to other inverter topologies [4].

    A number of papers discuss different topologies, performance and control strategies for cascaded multi-level inverters [5,6,7,8]. Khajehoddin et. al in [9] presents a control strategy to control cascaded multi-level converters in a multi-string configuration for single-phase grid connected systems. The performance of symmetrical and asymmetrical single-phase cascaded multi-level inverters with respect to harmonics content, number of switches and voltage stress across the switch with photovoltaic cell as its input source is simulated in [10]. A 9-level cascaded multi-level inverter with solar energy using PWM technique that provides a high switching frequency is presented in [11]. Even though these papers discuss the performance and control of cascaded multi-level inverters, no paper discusses their shading performance.

    In this paper, a cascaded multi-level inverter based PV system is presented and its shading performance is evaluated. In a PWM inverter based system, as a large number of cells are connected in series, shading a single cell causes the current in the string of cells to fall to the level of the shaded cell; thus reducing the generated PV power output [12]. In this proposed system, as each PV module (having lesser number of cells in series) generates a single voltage level in a step-wise output waveform, shading effects under partial shading conditions are expected to be much benign. Furthermore, as each string of cells is individually connected to an isolated H-bridge module, under fully shaded conditions the H-bridge can be controlled to by-pass the shaded module. The proposed system is simulated and a hardware prototype is implemented to validate the proposed system. Results are presented comparing the shading performance of the proposed system and a conventional PWM inverter system.

    2. Materials and Method

    Figure 1 shows the 13-level cascaded multi-level inverter based PV system used for this study. Each phase of the three-phase 13-level inverter consists of six H-bridge units connected in series.

    Figure 1. A single phase of the 13-level inverter.

    2.1. The operation of the 13-level inverter

    The principle of operation of the 13-level inverter is to synthesize the output voltage of each module to form a step-like ac voltage waveform across the output terminal. The ac voltage is produced by adding the output voltage of each module with different duty cycles. In general, with higher number of H-bridge modules in a single-phase structure, there will be more levels in the ac output voltage; thus producing an ac waveform closer to a sinusoidal wave. The number (M) of ac output voltage levels is given by M = 2N + 1, where N is the number of PV cells/modules. Then the output voltage, VO, of a single phase is given by,

    VO=VO1+VO2+.....+VO(N1)+VON (1)

    2.2. Switching pattern generator

    In an H-bridge there are two modes of operations depending on the pattern of the switching signals. This is explained using semiconductor switching devices S 11, S 12, S 13, and S 14 of the top most Hbridge in Figure 1.

    In the first mode, S 12 and S 14 are turned on to produce an output of +Vdc/6 and S 11 and S 13 are turned on to generate an output of -Vdc/6. The second mode gives an additional voltage level of 0 V: this occurs when the switch pairs S 12 and S 13, or S 11 and S 14, are turned on. The fundamental operation principle of the cascaded H-bridge multilevel inverter relies on the second mode of operation. In this mode, four switches in the H-bridge module is switched in four different sequences to generate a 3-level output voltage across output terminal of each H-bridge module as shown in Figure 2.

    Figure 2. Switching devices gate signal and 3-level output voltage waveform.

    2.2.1. Switching pattern of the 13-level inverter

    The output of the 13-level inverter is shown in Figure 3. This allows VO to vary between +Vdc and -Vdc in thirteen steps. This stair-case shape is obtained by switching individual bridges at different times corresponding to angles β1, β2, ...., and β6. Switching patterns are given in Table 1 and Table 2 for the positive half cycle and the negative half cycle respectively.

    Figure 3. 13-level H-Bridge output voltage.
    Table 1. Switching states for the 13-level inverter to produce positive voltages.
    Inverter output voltage (V)S 11 to S 61S 12S 22S 32S 42S 52S 62S 13 to S 63S 14 to S 64
    Vdc/6010000001
    2Vdc/6011000001
    3Vdc/6011100001
    4Vdc/6011110001
    5Vdc/6011111001
    6Vdc/6011111101
     | Show Table
    DownLoad: CSV
    Table 2. Switching states for the 13-level inverter to produce negative voltages.
    Inverter output voltage (V)S 11S 21S 31S 41S 51S 61S 12 to S 63S 13 to S 63S 14 to S 64
    0000000010
    -Vdc/6100000010
    -2Vdc/6110000010
    -3Vdc/6111000010
    -4Vdc/6111100010
    -5Vdc/6111110010
    -6Vdc/6111111010
     | Show Table
    DownLoad: CSV

    2.2.2. Selection of angles β1, β2, ...., β6

    The switching delay angles, β1, β2, ...., β6 shown in Figure 3 are obtained by minimising the sum of the squared error between the actual output of the converter and a reference sinusoidal wave. Due to the symmetry of the output waveform, only one quarter cycle was considered in the optimization. The objective function of this optimisation algorithm is to minimise the following function:

    T/4t=ot(Vmaxsin(2πft)Vdc66i=6a1(t))2 (2)
    subjected to: i:6,ai(t)<ai(t+t) where ai(t) is the state (1 when output is Vdc/6 and 0 when output is zero) of each H-bridge at time t.

    Using this optimisation, it was found that β1 = 5°, β2 = 15°, β3 = 25°, β4 = 36°, β5 = 49° and β6 = 67°.

    2.2.3. Partial shading performance balancing

    In the 13-level converter considered, the bridge switched on at the delay angle β1 will supply power from β1 to (π - β1). The PV module connected to this bridge will contribute more than any other PV module to form the stepwise ac waveform. On the other hand, the bridge switched at β6 will supply power from β1 to (π - β1) causing its PV module to contribute less than any other PV module when forming the stepwise ac waveform. In this operation, if a particular PV module is shaded, then it will always affect only a single step in the stepwise ac waveform. In order to circumvent the effect of shading on the output waveform, each bridge is switched on at different delay angle in each cycle. For example, the first bridge is switched on at β1 in the first cycle, at β2 in the second cycle, and at β6 in the sixth cycle. This equalises the contribution of all the PV modules over a period of six cycles on voltage waveform and the amount of energy supplied. The switching pulse generator achieves this by comparing the modulating signal with a stair-case carrier signal of period 6T with a six distinct voltage levels at sin(β1), sin(β2), sin(β3), sin(β4), sin(β5) and sin(β6) as shown in Figure 4. The width of each step of the carrier signal is equal to T. Figure 4 also illustrates how a switching signal is generated for S 13 (in Bridge 1). The 6-step stair-case carrier signal is compared to a sinusoidal modulating signal: when the modulating signal is greater than the carrier signal, the signal generator gives an output of 1; otherwise it gives an output of 0. To generate the switching signal to S 23 (in Bridge 2), the carrier signal in Figure 4 is shifted to the right by time T seconds, such that when the carrier signal of Bridge 1 is sin(β1), the carrier signal of Bridge 2 is sin(β2).

    Figure 4. Generation of switching signal for switch S 13 of Bridge 1 of the inverter.

    3. Results

    3.1. Hardware implementation of the single H-bridge module

    The structure of a single H-bridge module in the 13-level inverter is shown in Figure 5. It mainly consists of a separate PV module and four semiconductor switching devices. As the data were not available for the PV modules, its I-V characteristic was experimentally obtained and shown in Figure 6. The switching devices used were IRF840 MOSFETs. Other than a dc source and semiconductor switching devices, MOSFET driver ICs, Opto-couplers and dc-dc converters were used in this H-bridge module for signal isolation purposes.

    Figure 5. Single H-Bridge module.

    Figure 6. PV characteristic curve.

    An IR2110 MOSFET driver IC was used to produce necessary high-side and low-side gate signals for the H-bridge inverter. When cascading two H-Bridges, the gate signals provided to the second tier H-Bridge need to be boosted up further with respect to the ground. Since it is not practical to boost up gate signals of each H-Bridge, isolated gate signals were provided with respect to a reference point. To isolate the driver ICs, two dc-dc converters were used to provide isolated supply voltages (5 V and 15 V). Opto-couplers were used to isolate the signal level gate signals. The single H-bridge module was fabricated on a PCB as shown in Figure 7.

    Figure 7. H-Bridge module.

    The implemented H-bridge module was tested under different loading conditions where the dc link voltage was supplied from a PV module. Figure 8 shows the output voltage waveform of a single H-bridge inverter in an open circuit condition and in a loading condition (a 6 Ω resistor was connected to the output as a load) respectively.

    Figure 8. 3-level inverter output.

    3.2. The 13-level inverter

    The 13-level inverter was implemented as a laboratory prototype by cascading six H-bridge modules. It comprises of six H-bridge modules having twenty four MOSFET (IRF840) switches along with their driver circuits. An STM32F4 discovery board was used as the control environment to generate gate pulses for the inverter switches.

    The implemented 13-level inverter is shown in Figure 9. The inverter was tested under laboratory conditions and the setup is shown in Figure 10.

    Figure 9. Hardware prototype of 13-level inverter.

    Figure 10. Testing setup.

    Figure 11 shows the experimental output waveform and frequency spectrum of the output of the multi-level inverter under the open circuit condition. Figure 12 shows the experimental output waveform and frequency spectrum under loaded conditions. The load was maintained nearly at the maximum power point. Note that, in order to show the harmonic components properly, the spectrum is zoomed in: thus the fundamental component, which is 1 pu, is not properly displayed.

    Figure 11. Output voltage under open circuit condition.
    Figure 12. Output voltage under loaded condition.

    The peak of the voltage of inverter was 25.2 V at the open circuit condition, where the dc link (PV module) voltages of each H-bridge module were kept at 4.3 V.

    3.3. Harmonic performance of the 13-level inverter with shading

    Figure 13 and Figure 14 show the amplitude spectrum and output voltage waveform of the inverter at loading condition when one PV module is shaded and when two PV modules are shaded respectively. It was found that when two PV modules are shaded, the third harmonic component was high and the fifth harmonic component was low.

    Figure 13. One module is shaded at loading condition.
    Figure 14. Two modules are shaded at loading condition.

    The harmonic components that appear in the output voltage for different conditions are given in Table 3. When the amplitude spectrum of non-shaded condition is considered, it can be seen that the third harmonic component is low at loading condition than the open circuit condition. The amplitude spectrum of partially-shaded condition and fully-shaded condition of a single PV module has no reasonable variation.

    Table 3. Frequency analysis of multi-level inverter.
    ConditionFrequency
    Normalized amplitude
    Fundamental2nd3rd5th7th
    No loadNo shade10.0080.0530.0180.008
    1 module is shaded10.0080.0430.0240.018
    2 modules are shaded10.010.0380.0080.038
    Loaded (I = 40 mA)No shade10.0050.0280.0230.018
    1 module is shaded100.0480.0730.008
    2 modules are shaded10.0180.1060.0180.013
    1 module is partially shaded10.0580.050.0770.008
     | Show Table
    DownLoad: CSV

    4. Discussion

    4.1. Effect of shading: RMS value of the fundamental of the output voltage

    The significance of the proposed cascaded multi-level topology, in contrast to a PWM inverter (a single-phase single H-bridge unit with uni-polar PWM switching) was appreciated by examining the effect of shading on the fundamental of the output voltage value for both topologies. Using FFT analysis on the output voltages, the output voltage’s fundamental component was evaluated; the results are shown in Figure 15. Here, shaded cells are randomly chosen. It can be observed that the fundamental voltage is significantly reduced in the conventional method as the shading is increased from 0% to 100%. However, the difference in the fundamental voltage levels between the two methods for same shading conditions is slightly reduced as the number of shaded cells are increased.

    Figure 15. Effect of shading on the RMS value of the output voltage’s fundamental.

    4.2.1. Switching losses

    Consider the proposed modular approach with n number of levels. Therefore, the voltage across each bridge is Vdc/n; and the voltage across each semiconductor switch in blocking state (VT) is also Vdc/n. If the period of the fundamental is T1, according to Tables 1 and 2, the number of switches switched for a period of T1/2 is 2n.

    Number of switches switched for a period of T1=22n (3)

    Assume that the switching loss associated with a switch is a linear function of on-state current (kiIT) and blocking state voltage (kvVT) across the switch such that the energy loss (Esw) can be approximated as follows (see Figure 16);

    Esw=kvVT×kiIT×ktailIT (4)
    where, kv, ki and ktail are constants assumed to be only device dependent, IT is the constant output current of the PV system, ktail determines the rise and fall time of the current through the switch during turn-on and turn-off transients; hence, the term ktailIT signifies the duration of the switching transient.

    Figure 16. Turn-off switching transient: (a) ideal case, (b) approximated case.

    Now, the total switching loss in the cascaded multi-level topology during a fundamental cycle is calculated as;

    PlossSW,M=Esw4nT1 (5)

    Noting that VT = Vdc/n in eq. (4), using eqs. (4) and (5), we have;

    PlossSW,M=4k0VdcI2TT1 (6)
    where, ko is the simple product of the constants kv, ki and ktail.

    Now consider the PWM inverter; eq. (4) directly applies here as well. However, the total switching loss is defined as follows for this case;

    PlossSW,C=Esw4Tsw (7)
    where, Tsw is the switching frequency of the PWM inverter.

    Now using eqs. (4) and (7) and identifying that VT = Vdc;

    PlossSW,C=4k0VdcI2TTsw (8)

    Finally, getting the ratio of the switching losses between the two methods, we have (the subscripts M and C refers to multi-level and PWM approaches);

    PlossSW,CPlossSW,M=T1Tsw=fswf1=fm (9)
    where, fm is referred to as the frequency modulation ratio.

    With this result it is evident that the switching losses between the two approaches are ultimately defined by the frequency modulation ratio of the PWM approach. Therefore, in a qualitative sense, the number of levels in the modular approach has no effect on the ratio between the switching losses; however, it does have an effect on the absolute losses. As f1 is 50 Hz and fsw is usually selected around 1 - 2 kHz; fm is around 10-20. That is, the PWM inverter has 10-20 times more losses than that of the multi-level inverter.

    4.2.2. Conduction losses

    The steady state current through the switches in on-state for both approaches is the same and equal to IT. Therefore, at a glance it seems that the multi-level approach causes higher conduction losses due to the increased number of semiconductor switches. However, note that the blocking voltage (VT) required in the multi-level approach is 1/n th of that of the conventional approach: i.e. Vdc/n. This means that for the multi-level approach, semiconductor switches with lower blocking voltages can be chosen; which have the advantageous characteristic of having lower on-state resistances (for power MOSFETs) or on-state voltages (for IGBTs):

    PlossCON=I2T×Ronfor MOSFETSPlossCON=VCEon×ITfor IGBTs

    Therefore, the conduction losses associated with the switches would be actually lower than expected with the multi-level approach, provided that suitable semiconductor switches are chosen.

    4.3. Comparison of power utilization between the multi-level method and the conventional method

    With the conventional approach, with the aid of the intermediate dc-link and appropriate control, the complete PV unit is operated at the maximum power. However, the proposed method, with individual switching of each cell based upon angle β, the maximum power is not always extracted from each cell. Instantaneous utilization of this power depends upon the current operating position within the fundamental cycle (i.e. how many units are turned on) and depends upon the used load conditions as well: see Figure 17. For example, when β = 30° (three units ON) and RL = 40 Ω, the operation point is at point x marked on the plot. Therefore, as far as the zero shading condition is concerned, the conventional method implies better power utilization.

    Figure 17. Load-line and I-V curve intersections depending upon instantaneous number of units that are turned ON in the modular method (for a PV unit of: n = 36, Rp = 6.6 Ω, Rs = 0.005 Ω, Io = 6.00E - 10 A, Isc = 3.4 A).

    On the other hand, in practical applications there will always be the effect of shading. In such cases, the contrast between the power utilization between the two approaches is quite different. In the modular approach, few redundant PV units are kept in the system to be used under shading conditions while the shaded unit is bypassed: therefore, the power utilization is unaltered in the modular approach.

    However, the shading will affect the conventional approach due to the series connection of the PV units in the array. The effect of this on the two systems is illustrated in Figure 18. It is evident that the instantaneous power output of the conventional method deteriorates with the increasing number of shaded units in a series-connected array (due to the drop in the available total output voltage). Therefore, under shading conditions, the modular approach presents an appealing utilization of power: for instance, when 2 units are fully shaded, the modular approach offers an impressive advantage over the conventional method (see Figure 18).

    Figure 18. Comparison of power utilization between two approaches.

    5. Conclusion

    PV systems are now considered as a promising option for electric vehicles. As these vehicles are constantly subjected to shading, and since the shading on cells is dynamically changing, a shadingproof inverter is ideally suited for such applications.

    A development of a cascaded multi-level inverter based PV system is proposed for electric vehicle applications as a shading-proof inverter. The basic architecture and switching of the converter switches are described. A laboratory prototype of the proposed architecture was implemented using MOSFETs and harmonic performance under different shading conditions was evaluated. With a 13-level output, it was found that, except the third harmonic component, all the other voltage harmonics are below the values specified in the standards. It was also found that the third harmonic component increases with the shading on the cells/modules.

    The shading performance, losses and power utilization of the cascaded multi-level inverter were compared with that of a conventional PWM inverter. From the results, it was found that the proposed inverter shows better immunity for shading than a PWM inverter. As the ratio of the losses of a PWM inverter to the proposed inverter is equal to the frequency modulation index, the switching losses of the proposed inverter is 1/10 th to 1/20 th of that of a PWM inverter. Furthermore, the conduction losses, when compared to a PWM inverter, can be optimized by properly selecting the blocking voltage of the semiconductor switches used. Even though the power utilization is compromised at full insolation, the power utilization performance of the proposed inverter is superior under shading conditions, thus ideally suited for electric vehicle applications.

    The modular nature of the proposed inverter facilitates the cascading of more H-bridges with less cells/modules. This not only increases the number of steps in the synthesized sine wave, but also reduces the blocking voltage of the semiconductor switches. More levels will have a greater influence on the harmonic, shading, loss and power utilization performance of the proposed inverter.

    Acknowledgements

    The authors would like to thank the University of Peradeniya for the financial support through the grant RG/AF 2013/26/E.

    Conflict of Interest

    All authors declare no conflicts of interest in this paper.

    [1] Liou J, Kim ML, Heo WD, et al. (2005) STIM is a Ca2+ sensor essential for Ca2+-store-depletion-triggered Ca2+ influx. Curr Biol 15: 1235–1241. doi: 10.1016/j.cub.2005.05.055
    [2] Roos J, DiGregorio PJ, Yeromin AV, et al. (2005) STIM1, an essential and conserved component of store-operated Ca2+ channel function. J Cell Biol 169: 435–45. doi: 10.1083/jcb.200502019
    [3] Feske S, Gwack Y, Prakriya M, et al. (2006) A mutation in Orai1 causes immune deficiency by abrogating CRAC channel function. Nature 441: 179–185. doi: 10.1038/nature04702
    [4] Vig M, Peinelt C, Beck A, et al. (2006) CRACM1 is a plasma membrane protein essential for store-operated Ca2+ entry. Science 312: 1220–1223. doi: 10.1126/science.1127883
    [5] Zhang SL, Yeromin AV, Zhang XH-F, et al. (2006) Genome-wide RNAi screen of Ca(2+) influx identifies genes that regulate Ca(2+) release-activated Ca(2+) channel activity. Proc Natl Acad Sci U S A 103: 9357–9362. doi: 10.1073/pnas.0603161103
    [6] Liou J, Fivaz M, Inoue T, et al. (2007) Live-cell imaging reveals sequential oligomerization and local plasma membrane targeting of stromal interaction molecule 1 after Ca2+ store depletion. Proc Natl Acad Sci U S A 104: 9301–9306. doi: 10.1073/pnas.0702866104
    [7] Hou X, Pedi L, Diver MM, et al. (2012) Crystal structure of the calcium release-activated calcium channel Orai. Science 338: 1308–1313. doi: 10.1126/science.1228757
    [8] Korzeniowski MK, Manjarrés IM, Varnai P, et al. (2010) Activation of STIM1-Orai1 involves an intramolecular switching mechanism. Sci Sign 3: ra82.
    [9] Muik M, Fahrner M, Schindl R, et al. (2011) STIM1 couples to ORAI1 via an intramolecular transition into an extended conformation. EMBO J 30: 1678–1689. doi: 10.1038/emboj.2011.79
    [10] Zhou Y, Srinivasan P, Razavi S, et al. (2013) Initial activation of STIM1, the regulator of store-operated calcium entry. Nat Struct Mol Biol 20: 973–981. doi: 10.1038/nsmb.2625
    [11] Stathopulos PB, Schindl R, Fahrner M, et al. (2013) STIM1/Orai1 coiled-coil interplay in the regulation of store-operated calcium entry. Nat Commun 4: 2963.
    [12] Kawasaki T, Lange I, Feske S (2009) A minimal regulatory domain in the C terminus of STIM1 binds to and activates ORAI1 CRAC channels. Biochem Biophys Res Comm 385: 49–54. doi: 10.1016/j.bbrc.2009.05.020
    [13] Yuan JP, Zeng W, Dorwart MR, et al. (2009) SOAR and the polybasic STIM1 domains gate and regulate Orai channels. Nat Cell Biol 11: 337–343. doi: 10.1038/ncb1842
    [14] Park CY, Hoover PJ, Mullins FM, et al. (2009) STIM1 Clusters and Activates CRAC Channels via Direct Binding of a Cytosolic Domain to Orai1. Cell 136: 876–890. doi: 10.1016/j.cell.2009.02.014
    [15] Yang X, Jin H, Cai X, et al. (2012) Structural and mechanistic insights into the activation of Stromal interaction molecule 1 (STIM1). Proc Natl Acad Sci U S A 109: 5657–5662. doi: 10.1073/pnas.1118947109
    [16] Hull JJ, Lee JM, Kajigaya R, et al. (2009) Bombyx mori homologs of STIM1 and Orai1 are essential components of the signal transduction cascade that regulates sex pheromone production. J Biol Chem 284: 31200–31213. doi: 10.1074/jbc.M109.044198
    [17] Calloway N, Holowka DA, Baird BA (2010) A Basic Sequence in STIM1 Promotes Ca2+ Influx by Interacting with the C-Terminal Acidic Coiled Coil of Orai1. Biochemistry 49: 1067–1071. doi: 10.1021/bi901936q
    [18] Barr VA, Bernot KM, Srikanth S, et al. (2008) Dynamic movement of the calcium sensor STIM1 and the calcium channel Orai1 in activated T-cells: puncta and distal caps. Mol Biol Cell 19: 2802–2817. doi: 10.1091/mbc.E08-02-0146
    [19] Varnai P, Tóth B, Tóth DJ, et al. (2007) Visualization and manipulation of plasma membrane-endoplasmic reticulum contact sites indicates the presence of additional molecular components within the STIM1-Orai1 Complex. J Biol Chem 282: 29678–90. doi: 10.1074/jbc.M704339200
    [20] Holowka D, Korzeniowski MK, Bryant KL, et al. (2014) Polyunsaturated fatty acids inhibit stimulated coupling between the ER Ca(2+) sensor STIM1 and the Ca(2+) channel protein Orai1 in a process that correlates with inhibition of stimulated STIM1 oligomerization. Biochim Biophys Acta 1841: 1210–1216. doi: 10.1016/j.bbalip.2014.04.006
    [21] Wissenbach U, Philipp SE, Gross SA, et al. (2007) Primary structure, chromosomal localization and expression in immune cells of the murine ORAI and STIM genes. Cell Calcium 42: 439–446. doi: 10.1016/j.ceca.2007.05.014
    [22] Wu MM, Covington ED, Lewis RS (2014) Single-molecule analysis of diffusion and trapping of STIM1 and Orai1 at ER-plasma membrane junctions. Mol Biol Cell 25: 3672-3685.
    [23] Gwozdz T, Dutko-Gwozdz J, Zarayskiy V, et al. (2008) How strict is the correlation between STIM1 and Orai1 expression, puncta formation, and ICRAC activation? Am J Physiol Cell Physiol 295: C1133–1140. doi: 10.1152/ajpcell.00306.2008
    [24] Varnai P, Hunyady L, Balla T (2009) STIM and Orai: the long-awaited constituents of store-operated calcium entry. Trends Pharmacol Sci 30: 118–128. doi: 10.1016/j.tips.2008.11.005
    [25] Perni S, Dynes JL, Yeromin AV, et al. (2015) Nanoscale patterning of STIM1 and Orai1 during store-operated Ca2+ entry. Proc Natl Acad Sci U S A 112: E5533-42.
    [26] Jha A, Ahuja M, Maléth J, et al. (2013) The STIM1 CTID domain determines access of SARAF to SOAR to regulate Orai1 channel function. J Cell Biol 202: 71–79. doi: 10.1083/jcb.201301148
    [27] Muik M, Frischauf I, Derler I, et al. (2008) Dynamic coupling of the putative coiled-coil domain of ORAI1 with STIM1 mediates ORAI1 channel activation. J Biol Chem 283: 8014–8022. doi: 10.1074/jbc.M708898200
    [28] Covington ED, Wu MM, Lewis RS (2010) Essential role for the CRAC activation domain in store-dependent oligomerization of STIM1. Mol Biol Cell 21: 1897–1907. doi: 10.1091/mbc.E10-02-0145
    [29] Calloway N, Vig M, Kinet J, et al. (2009) Molecular clustering of STIM1 with Orai1/CRACM1 at the plasma membrane depends dynamically on depletion of Ca2+ stores and on electrostatic interactions. Mol Biol Cell 20: 389–399. doi: 10.1091/mbc.E07-11-1132
    [30] Stathopulos PB, Li G-Y, Plevin MJ, et al. (2006) Stored Ca2+ depletion-induced oligomerization of stromal interaction molecule 1 (STIM1) via the EF-SAM region: An initiation mechanism for capacitive Ca2+ entry. J Biol Chem 281: 35855–35862. doi: 10.1074/jbc.M608247200
    [31] Muik M, Fahrner M, Derler I, et al. (2009) A Cytosolic Homomerization and a Modulatory Domain within STIM1 C Terminus Determine Coupling to ORAI1 Channels. J Biol Chem 284: 8421–8426. doi: 10.1074/jbc.C800229200
    [32] Fahrner M, Muik M, Schindl R, et al. (2014) A coiled-coil clamp controls both conformation and clustering of stromal interaction molecule 1 (STIM1). J Biol Chem 289: 33231-33244. doi: 10.1074/jbc.M114.610022
    [33] Shim AH-R, Tirado-Lee L, Prakriya M (2015) Structural and functional mechanisms of CRAC channel regulation. J Mol Biol 1427: 77-93.
    [34] Mullins FM, Park CY, Dolmetsch RE, et al. (2009) STIM1 and calmodulin interact with Orai1 to induce Ca2+-dependent inactivation of CRAC channels. Proc Natl Acad Sci U S A 106: 15495–15500. doi: 10.1073/pnas.0906781106
    [35] Derler I, Fahrner M, Muik M, et al. (2009) A Ca2(+ )release-activated Ca2(+) (CRAC) modulatory domain (CMD) within STIM1 mediates fast Ca2(+)-dependent inactivation of ORAI1 channels. J Biol Chem 284: 24933–2498. doi: 10.1074/jbc.C109.024083
    [36] Lee KP, Yuan JP, Zeng W, et al. (2009) Molecular determinants of fast Ca2+-dependent inactivation and gating of the Orai channels. Proc Natl Acad Sci U S A 106: 14687–14692. doi: 10.1073/pnas.0904664106
  • This article has been cited by:

    1. Yash Shukla, Zaid Ahmad Khan, Khan Mohammad, M Saad Bin Arif, 2024, Comparative Analysis of Selected Reduced Switch Count NPC Multilevel Inverter Topologies for Solar PV Applications, 979-8-3315-4158-3, 589, 10.1109/I3CEET61722.2024.10993910
  • Reader Comments
  • © 2016 the Author(s), licensee AIMS Press. This is an open access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0)
通讯作者: 陈斌, bchen63@163.com
  • 1. 

    沈阳化工大学材料科学与工程学院 沈阳 110142

  1. 本站搜索
  2. 百度学术搜索
  3. 万方数据库搜索
  4. CNKI搜索

Metrics

Article views(6649) PDF downloads(1461) Cited by(7)

Article outline

Figures and Tables

Figures(5)  /  Tables(1)

/

DownLoad:  Full-Size Img  PowerPoint
Return
Return

Catalog