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2016-01-21

Usb Embedded Device Driver

Deployed in over half of all current embedded USB modems, USBware Device is the world s most widely used USB device stack. This modular USB protocol software.

PDF Version

Ways to Use USB in Embedded Systems

by Yingbo Hu, R D Embedded Software Engineer

and Ralph Moore, President of Micro Digital

Universal Serial Bus USB is a connectivity specification that provides ease of use, expandability, and good performance for the end user. It is one of the most successful interconnects in computer history. Originally released in 1995 for PCs, it now is expanding into use by embedded systems and is replacing older interfaces such as serial and parallel interfaces as the preferred communication link. This article has been written as a tutorial on the many ways that USB can be employed in embedded systems.

Introduction

USB is not a peer-to-peer protocol like Ethernet. One USB device, called the USB host, acts as the master. Other USB devices, called USB devices or USB peripherals, act as slaves. The host initiates all bus transfers. Up to 127 USB devices can be connected to one USB host via up to 6 layers of cascaded hubs. For embedded systems, it is very unusual to have more than one hub. In most cases, one USB device connects directly to one USB host with no hub.

A USB host requires a USB host controller and USB host software. The latter is layered from the bottom up as follows: 1 USB host controller driver, 2 USB host stack, and 3 USB class driver. The first layer controls the USB host controller – i.e. it reads and writes registers in the controller and it transfers data. The second layer implements the USB protocol and thus controls connected USB devices. The third layer is device-aware and communicates with and controls the actual device e.g. disk drive, HID human interface device, CDC communication device, etc. One USB host stack can support multiple class drivers, simultaneously. In an embedded system there usually is only one USB host controller.

A USB device requires a USB device controller and USB device software. The latter is layered from the bottom up as follows: 1 USB device controller driver, 2 USB device stack, and 3 USB function driver. The first layer controls the USB device controller – i.e. it reads and writes registers in the controller and it transfers data. The second layer implements the USB protocol and thus communicates with the USB host stack. The third layer communicates with the class driver in the host and provides the actual device control. It makes the embedded unit look like a USB disk drive, HID, serial device, etc. One USB device stack can support more than one function driver simultaneously, through the composite device framework.

A nice feature of USB is it is plug and play, which means that a USB device will be automatically recognized shortly after being connected to a host. Also, cabling is simple: There is an A receptacle/plug pair for the host end and a B receptacle/plug pair for the device end. All hosts and devices adhere to this standard, except On The Go OTG devices, which are designed for but not widely used yet in embedded systems.

Example Configurations

Following are examples of how USB can be utilized in the embedded system, for both host and device. Where performance information is given, a medium performance processor is assumed to be a 50-80 MHz ARM7 or ColdFire.

Figure 1: PC to Device via USB Serial

Most new PCs and laptops do not provide serial or parallel ports; they have been replaced with USB ports. Hence, connecting a PC to embedded device via its RS232 port is no longer possible. As part of their USB host stacks, popular PC operating systems Windows, Mac OS, and Linux include Communication Class Drivers CDC. As shown in Figure 1, if the embedded device has a Serial/CDC Function Driver then it will look like a serial device to the PC. When it is plugged in, it will be recognized by the PC OS as a serial device, and it will be automatically assigned a COM port number. Then, terminal emulators and other serial applications can communicate with the embedded device without any modification. Hence, we are back to where we started, which, in this case, is a good thing. This use of USB is particularly good for control and transferring serial data. Transfer rates of 800 KB/sec are feasible at full speed and 2500 KB/sec at high speed for medium speed embedded processors.

Figure 2: PC to Device via USB Disk

Another method to connect a PC or laptop to an embedded device is for the embedded device to emulate a USB disk drive. Popular PC operating systems have built-in USB mass storage class drivers that interface their file systems to the USB host stack, as shown on the left side of Figure 2. Adding a mass storage function driver to the embedded device enables it to look like a USB disk drive to the PC. Also shown, as an example, is how a resident flash memory can be accessed as a flash disk via the USB function driver connected to its flash driver. Any other type of storage media could be connected, instead, via its own driver.

When the embedded device is plugged into a PC, it is recognized as a disk drive and automatically assigned a drive letter. Thereafter, files can be dragged and dropped to and from the embedded device as though it were a disk drive. In this example, a PC application could read and write the files on the flash disk. Note that the embedded application uses a local file system, as shown in Figure 2, to access the flash disk, itself. This file system must, of course, be Windows-compatible. An important concept to understand is that within the PC, the PC s file system is used and the embedded device merely looks like another disk drive to it. This use of USB would be particularly good for uploading acquired data files or downloading new versions of code files.

Figure 3: Web Server Access via USB RNDIS

RNDIS Remote Network Driver Interface Specification permits emulating Ethernet over USB. It is not part of the USB specification, but some popular PC operating systems, such as Windows and Linux, support it. As shown in Figure 3, adding an RNDIS function driver to an embedded device allows interfacing its USB device stack to its TCP/IP stack that, in turn, connects to its web server. When the embedded device is plugged into a PC, its browser can connect to the web server in the embedded device. Hence, it is possible to use a browser to access an embedded device s web server, even when there is no Ethernet connection or it is difficult to access. This can be convenient for field troubleshooting or configuration using a laptop. The same information accessed via the network to which the embedded device is connected, can be accessed via USB.

Figure 4: USB Audio Device with MIDI

The USB Audio Class is defined such that if a USB device follows the specification, a PC host will treat it as a sound card. As shown in Figure 4, this is accomplished by adding an audio function driver to the embedded device. This function driver provides both an audio interface and a MIDI Musical Instrument Digital Interface interface to a sound manager, which in turn can connect to a microphone, speaker, or other audio gear, such as a musical instrument. Thus audio and MIDI streams can be transferred over a USB link rather than using audio or MIDI cables. This uses the USB isochronous mode of data transfer.

Figure 5: USB Multi-Port Serial Device with UART and Other Connections

In Figure 1 we examined the case of one serial channel over a USB connection. However it is actually possible to run multiple, independent serial channels over one USB connection. This is practical because of the higher speed of USB, as noted in the Figure 1 discussion. Figure 5 shows the block diagram. The CDC ACM class driver in the PC may not be the native driver that comes with the PC OS. A special driver may need to be installed. This driver presents multiple virtual COM ports to the PC application and it multiplexes the corresponding serial channels over the USB connection.

In the embedded device, the USB CDC function driver de-multiplexes the serial channels. Note that, in this example, one channel goes to an application task, which might return certain internal information, and the other two serial channels connect to actual UARTs. The application in the PC can communicate with physical devices, such as modem, bar code reader, printer, etc. connected to the UARTs as though they were connected directly to serial ports on the PC which we know does not actually have serial ports, anymore. The high throughput of USB makes multiple channels feasible. For example, with a medium performance processor and full speed USB, a total bandwidth of 200 KB/sec is achievable. This would support 15 115.2 Kbaud channels, with capacity left over.

Figure 6: USB Composite Devices

It is actually possible for one USB device to look like multiple USB devices to a USB host, simultaneously. This is made possible by the USB Composite Device Framework, as shown in Figure 6. The USB host PC in this example will recognize each USB device within the embedded device and load its corresponding class driver. In Figure 6 the device looks like a USB disk and a serial port. Note that both function drivers are present. This example is a fairly common case that is supported by PC OSs. Many possible combinations are not supported by PC OSs. This particular one would support an application in the PC transferring files and another application allowing an operator to control or configure the embedded device.

Figure 7: USB Thumb Drive Support

Figure 7 shows how an embedded device can access a USB thumb drive also called a USB memory stick. A mass storage class driver fits between the USB host stack and the local file system in the embedded device. It creates the usual read/write logical address API expected of media drivers. Naturally the file system must be OS-compatible in order to exchange thumb drives with a PC. Thumb drives are commonly used to transfer data from embedded devices to PCs or to update firmware or configuration settings and tables in embedded devices.

Figure 8: USB to Ethernet

If an embedded device already has USB host capability, it is possible to connect it to a local Ethernet network using a USB to Ethernet adapter. This can be done without adding hardware to the embedded device. Figure 8 shows the software required to do the job. The USB to Ethernet class driver interfaces the USB host stack to a TCP/IP stack in the embedded device. This allows some units to connect to a LAN without the expense of adding Ethernet connectivity to all units. It thus allows greater flexibility in meeting customer requirements and saves redesign.

Figure 9: USB to WiFi

Wireless communication is becoming more and more popular. Several vendors are providing USB to WiFi 802.11 chipsets which enable systems having USB host ports to add wireless connectivity. These chipsets are commercially available in what are called WiFi dongles or WiFi keys and are generally inexpensive. Figure 9 shows the software needed in the embedded device. The 802.11 Media Access Controller provides an Ethernet-like interface to the local TCP/IP stack and controls the 802.11 controller in the WiFi chipset. The WiFi chipset driver controls the USB interface in the chipset. For security, 802.11 MAC also provides WEP Wired Equivalency Privacy or WPA WiFi Protected Access.

Like the USB to Ethernet capability shown in Figure 8, this feature can be added to only some embedded units, as required. It is very useful in the field when a wired connection is not available or too expensive. Transfer rates of 200 KB/sec are typical for medium performance processors.

Figure 10: USB Modem Access

If an embedded device already has USB host capability, adding a USB CDC-ACM class driver can be used to add dialup modem capability to the system. This is shown in Figure 10. No hardware change is necessary. USB modems are available that connect to phone lines in the PSTN system. Some cell phones that have a USB modem interface can be used to connect wirelessly. Hence, this can be a flexible option for connecting to an embedded device. It is especially useful when the dialup communication is only needed occasionally. Also, one USB modem can be shared between multiple devices to save the cost of adding modem hardware. Dialup connections are very useful for long distance communication.

Figure 11: Connection to Multiple Sensors and Actuators

Figure 11 shows how an external hub can be used to connect an embedded control unit to multiple sensors and actuators. In this diagram, it is assumed that the sensors are serial devices. They are handled by a serial class driver. The actuators are assumed to be custom devices requiring custom class drivers. A well-structured USB host stack permits easily adding custom class drivers. Standard USB peripherals, such as printers and bar-code readers could also be added. They would be supported by standard class drivers. For example a keyboard or joystick would be supported by an HID class driver. A gaming machine is a good example of a unit incorporating custom sensors and actuators and standard USB peripherals.

Conclusion

USB usage in embedded systems so far has been largely centered on dealing with the loss of serial and parallel ports on PCs and laptops, the loss of parallel interface printers, and with capitalizing on the low cost and convenience of USB thumb drives for transporting information. However, as we hope this article has shown, USB offers many other capabilities that are available to solve other problems in the embedded space. We expect to see these uses grow in the future.

Micro Digital s software supports all of the configurations described in this article.

Yingbo Hu is the lead developer of smxUSBD, smxUSBH, and smxWiFi, as well as smxFS and smxFFS products of Micro Digital Inc. He specializes in writing device drivers for embedded peripherals, such as USB and flash, and he has 11 years experience developing real-time embedded software. He graduated with a Masters Degree from Beijing University of Aeronautics and Astronautics.

Ralph Moore is the current President and founder of Micro Digital, Inc. He graduated from Caltech with a Bachelors degree in physics long before the first embedded system or even the first integrated circuit. He has watched our industry develop with amazement. Mr. Moore is also the architect of the smx real time multitasking kernel.

For More Information

smx is a registered trademark of Micro Digital Inc. smx product names are trademarks of Micro Digital Inc.

2/12/09

back to White Papers page.

Ways to Use USB in Embedded Systems

SmxUSBD is a USB device stack for ARM, ColdFire, and other embedded systems. Mass storage, serial, audio, RNDIS. ISPxxxx and SoC controller drivers. No royalty.

2008 Microchip Technology Inc. DS01164A-page 3 AN1164 SOURCE FILE ORGANIZATION The CDC serial device USB stack contains the following source and.

Thesycon develop universal drivers, driver templates and development toolkits for Windows operating systems. It includes software toolkit for USB. The toolkit.

Bill Gatliff-December 17, 2002 Linux-based USB Devices The Universal Serial Bus is a useful communications interface and more popular than ever.

Thesycon s USB device software stack implements the core functionality of a USB device and provides device class specific protocols. It enables developers to easily.

AN1144 DS01144A-page 2 2008 Microchip Technology Inc. THE HID CLIENT DRIVER Architecture of HID Client Driver Applications do not interface directly with the USB HID.

Provides WinDriver and KernelDriver device driver development toolkits and HotSwap solutions for Windows, Linux, Solaris.

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Lagarith Lossless Video Codec 1.3.27

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LAV CUVID Decoder 0.13

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LAV Filters 0.67

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Ligos Indeo Codec 5.11

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LZOcodec 0.4

OBSOLETE Updated: 6 Nov 2006 License: Freeware Size: 44KB Downloads: 8676

LZOcodec enables you to perform lossless video compression in 3 steps: converting RGB information to YUY2, creating a compression algorithm and eventually using the LZO algorithm.

madVR 0.89.19

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MagicYUV 1.2

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MainConcept DV Codec 2.4.16

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MainConcept Motion JPEG Codec 3.2.4

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Matrox Codecs 2.0

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Midvid JPEG Video Codec 1.0

OBSOLETE Updated: 31 Dec 2004 License: Freeware Size: 175KB Downloads: 21153

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Midvid Mideo Codec 1.06 beta

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Midvid MV3 Video Codec 1.0

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MindVid 1.0 Beta 1

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Moonlight Scalar DirectShow Filter 1.03

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OBSOLETE Updated: 28 Sep 2004 License: Freeware Size: 50KB Downloads: 12095

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FRESH Updated: 10 Dec 2015 License: Freeware Size: 603KB Downloads: 415161

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mpegable AVC 0.10

OBSOLETE Updated: 17 Sep 2004 License: Freeware Size: 1.1MB Downloads: 224874

mpegable AVC was designed to enable video compression with all applications based on VfW, such as VirtualDub, Windows Media Player, FlaskMPEG, etc.

mpegable DS decoder 2.1.4

OBSOLETE Updated: 17 Sep 2004 License: Freeware Size: 1.25MB Downloads: 370430

The mpegable DS decoder enables you to decode MPEG-4 and 3GPP audio and video using DirectShow-based applications, such as Windows Media Player 6.x and above.

MPV Decoder 1.4.6.1043

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MPV Decoder is a filter which allows you to decode MPV files. MPV is an MPEG-1 or MPEG-2 video which doesn t contain any audio streams.

MSU Lossless Video Codec 0.6.0

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MSU Screen Capture Lossless Codec 1.2

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NRW Codec 1.6

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nVIDIA PureVideo Decoder 1.02-223

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Ogg DirectShow Filters 0.9.9.6

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OLYMPUS RAW Codec 1.3

OBSOLETE Updated: 9 Jul 2009 License: Freeware Size: 5.1MB Downloads: 9231

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Panasonic DV Codec

OBSOLETE Updated: 8 Apr 2004 License: Freeware Size: 75KB Downloads: 334741

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Pegasus SVM Video Decoder 3.0

OBSOLETE Updated: 16 Jun 2005 License: Freeware Size: 393KB Downloads: 21043

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Pinnacle DV25 Codec 1.5.0.9

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Pinnacle MPEG Realtime Codec 2.0

OBSOLETE Updated: 2 Feb 2005 License: Freeware Size: 315KB Downloads: 55918

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PixiShow 2005-08-27

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PIXL Codec Pinnacle PCTV

OBSOLETE Updated: 30 Aug 2004 License: Freeware Size: 27KB Downloads: 9780

PIXL Codec, also known as Pinnacle PCTV, is an alias for VIXL, a JPEG format used in Zoran chipsets.

PlayFLV 1.00

OBSOLETE Updated: 2 Nov 2008 License: Freeware Size: 4.73MB Downloads: 113777

PlayFLV is a powerful tool making it possible to play FLV files in Windows Media Player.

RadLight PVA DirectShow filter 1.0

OBSOLETE Updated: 6 Feb 2005 License: Freeware Size: 244KB Downloads: 8601

The RadLight PVA DirectShow filter enables you to play PVA format files in RadLight 4 and in any other media player based on DirectShow.

REALmagic MPEG-4 Video Codec 1.1

OBSOLETE Updated: 9 Aug 2004 License: Freeware Size: 400KB Downloads: 56935

With the REALmagic MPEG-4 Video Codec from Sigma Design, you can create high-quality MPEG-4 video bitstreams which are ISO-compatible.

ReClock DirectShow Filter 1.8.8.5

Updated: 15 Apr 2015 License: Freeware Size: 1.1MB Downloads: 179463

ReClock DirectShow Filter helps you to avoid discontinuous playback of AVI and MPEG files, including those situations when you connect your PC to a TV set.

River Past QuickTime DirectShow Filter 1.0.16

OBSOLETE Updated: 5 May 2011 License: Freeware Size: 349KB Downloads: 3190

By installing River Past QuickTime DirectShow Filter to your computer, you will easily play QuickTime and MPEG4 files using Windows Media Player.

Screen Capture DirectShow Filter 2.0.3

Updated: 26 Jun 2015 License: Shareware Size: 2.9MB Downloads: 6444

The MediaLooks Screen Capture DirectShow Filter allows you to capture images from your computer screen while enjoying a wide range of features.

SIF1 Video Codec 1.20

OBSOLETE Updated: 13 Jul 2011 License: Freeware Size: 1MB Downloads: 8572

SIF1 Video Codec is a flexible application which can be used with any software in order to edit and view video files.

SolveigMM Subtitler 1.0.812.20

OBSOLETE Updated: 12 Feb 2011 License: Freeware Size: 68KB Downloads: 1281

SolveigMM Subtitler is a DirectShow filter which allows you to overlay Unicode text that is read from set SRT file to a specified position on raw video in RGB24 format.

Sony RAW Codec 2.0

OBSOLETE Updated: 27 Mar 2009 License: Freeware Size: 7.4MB Downloads: 19901

Sony RAW Codec enables you to work with. ARW. SRF and. SR2 files.

Valid FOURCC Codes blue FOURCC - decompression only / green - formats Notes Universal: LAV Filters v.0.63 32-bit 64-bit MKV WebM AVI MP4 MOV TS MPEG.

Confirmed as working. I edited a bunch of video using Canopus hardware which I ve long since stopped using. I installed this codec on Win 7 64 bit and can watch all the old avi files no problem.

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2016-01-21

Umax Astra 2000u Driver Xp

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2016-01-21

Activation Postsynaptic Membrane

Membrane potential also transmembrane potential or membrane voltage is the difference in electric potential between the interior and the exterior of a biological cell.

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An inhibitory postsynaptic potential IPSP is a kind of synaptic potential that makes a postsynaptic neuron less likely to generate an action potential. 1 The opposite of an inhibitory postsynaptic potential is an excitatory postsynaptic potential EPSP, which is a synaptic potential that makes a postsynaptic neuron more likely to generate an action potential. They can take place at all chemical synapses, which use the secretion of neurotransmitters to create cell to cell signalling. Inhibitory presynaptic neurons release neurotransmitters that then bind to the postsynaptic receptors; this induces a postsynaptic conductance change as ion channels open or close. An electric current that changes the postsynaptic membrane potential to create a more negative postsynaptic potential is generated. Depolarization can also occur due to an IPSP if the reverse potential is between the resting threshold and the action potential threshold. Another way to look at inhibitory postsynaptic potentials is that they are also a chloride conductance change in the neuronal cell because it decreases the driving force. 2 Microelectrodes can be used to measure postsynaptic potentials at either excitatory or inhibitory synapses.

In general, a postsynaptic potential is dependent on the type and combination of receptor channel, reverse potential of the postsynaptic potential, action potential threshold voltage, ionic permeability of the ion channel, as well as the concentrations of the ions in and out of the cell; this determines if it is excitatory or inhibitory. IPSPs always want to keep the membrane potential more negative than the action potential threshold and can be seen as a transient hyperpolarization. 3 EPSPs and IPSPs compete with each other at numerous synapses of a neuron; this determines whether or not the action potential at the presynaptic terminal will regenerate at the postsynaptic membrane. Some common neurotransmitters involved in IPSPs are GABA and glycine.

Flowchart describing how an inhibitory postsynaptic potential works from neurotransmitter release to summation

Contents

1 Components

1.1 Types

1.2 Factors

2 Inhibitory Molecules

3 Inhibitory Receptors

3.1 Ionotropic receptors

3.2 Metabotropic receptors

4 Significance

5 Studies

6 See also

7 References

Components edit

Types edit

This system 1 IPSPs can be temporally summed with subthreshold or suprathreshold EPSPs to reduce the amplitude of the resultant postsynaptic potential. Equivalent EPSPs positive and IPSPs negative can cancel each other out when summed. The balance between EPSPs and IPSPs is very important in the integration of electrical information produced by inhibitory and excitatory synapses.

Graph displaying an EPSP, an IPSP, and the summation of an EPSP and an IPSP. When the two are summed together the potential is still below the action potential threshold.

Factors edit

The size of the neuron can also affect the inhibitory postsynaptic potential. Simple temporal summation of postsynaptic potentials occurs in smaller neurons, whereas in larger neurons larger numbers of synapses and ionotropic receptors as well as a longer distance from the synapse to the soma enables the prolongation of interactions between neurons.

Inhibitory Molecules edit

GABA is a very common neurotransmitter used in IPSPs in the adult mammalian brain and retina. 1 4 GABA receptors are pentamers most commonly composed of three different subunits α, β, γ, although several other subunits δ,ε, θ, π, ρ and conformations exist. The open channels are selectively permeable to chloride or potassium ions depending on the type of receptor and allow these ions to pass through the membrane. If the electrochemical potential of the ion is more negative than that of the action potential threshold then the resultant conductance change that occurs due to the binding of GABA to its receptors keeps the postsynaptic potential more negative than the threshold and decreases the probability of the postsynaptic neuron completing an action potential. Glycine molecules and receptors work much in the same way in the spinal cord, brain, and retina.

Inhibitory Receptors edit

There are two types of inhibitory receptors:

Ionotropic receptors edit

Ionotropic receptors also known as ligand-gated ion channels play an important role in inhibitory postsynaptic potentials. 1 A neurotransmitter binds to the extracellular site and opens the ion channel that is made up of a membrane-spanning domain that allows ions to flow across the membrane inside the postsynaptic cell. This type of receptor produces very fast postsynaptic actions within a couple of milliseconds of the presynaptic terminal receiving an action potential. These channels influence the amplitude and time-course of postsynaptic potentials as a whole. Ionotropic GABA receptors are used in binding for various drugs such as barbiturates Phenobarbital, pentobarbital, steroids, and picrotoxin. Benzodiazepines Valium bind to the α and δ subunits of GABA receptors in order to improve GABAergic signaling. Alcohol also modulates ionotropic GABA receptors.

Metabotropic receptors edit

Metabotropic receptors, or G-protein-coupled receptors, do not use ion channels in their structure; they, instead, consist of an extracellular domain that binds to a neurotransmitter and an intracellular domain that binds to G-protein. 1 This begins the activation of the G-protein, which then releases itself from the receptor and interacts with ion channels and other proteins to open or close ion channels through intracellular messengers. They produce slow postsynaptic responses from milliseconds to minutes and can be activated in conjunction with ionotropic receptors to create both fast and slow postsynaptic potentials at one particular synapse. Metabotropic GABA receptors, heterodimers of R1 and R2 subunits, use potassium channels instead of chloride. They can also block calcium ion channels in order to hyperpolarize postsynaptic cells.

Significance edit

There are many applications of inhibitory postsynaptic potentials to the real world. Drugs that affect the actions of the neurotransmitter can treat neurological and psychological disorders through different combinations of types of receptors, G-proteins, and ion channels in postsynaptic neurons.

For example, studies researching opioid receptor-mediated receptor desensitizing and trafficking in the locus cereleus of the brain are being performed. When a high concentration of agonist is applied for an extended amount of time fifteen minutes or more, hyperpolarization peaks and then decreases. This is significant because it is a prelude to tolerance; the more opioids one needs for pain the greater the tolerance of the patient. These studies are important because it helps us to learn more about how we deal with pain and our responses to various substances that help treat pain. By studying our tolerance to pain, we can develop more efficient medications for pain treatment. 5

In addition, research is being performed in the field of dopamine neurons in the ventral tegmental area, which deals with reward, and the substantia nigra, which is involved with movement and motivation. Metabotropic responses occur in dopamine neurons through the regulation of the excitability of cells. Opioids inhibit GABA release; this decreases the amount of inhibition and allows them to fire spontaneously. Morphine and opioids relate to inhibitory postsynaptic potentials because they induce disinhibition in dopamine neurons. 5

IPSPs can also be used to study the input-output characteristics of an inhibitory forebrain synapse used to further study learned behavior, to be specific song learning in birds in a study performed at the University of Washington. 6 Poisson trains of unitary IPSPs were induced at a high frequency to reproduce postsynaptic spiking in the medial portion of the dorsalateral thalamic nucleus without any extra excitatory inputs. This shows an excess of thalamic GABAergic activation. This is important because spiking timing is needed for proper sound localization in the ascending auditory pathways. Songbirds use GABAergic calyceal synaptic terminals and a calcyx-like synapse such that each cell in the dorsalateral thalamic nucleus receives at most two axon terminals from the basal ganglia to create large postsynaptic currents.

Inhibitory postsynaptic potentials are also used to study the basal ganglia of amphibians to see how motor function is modulated through its inhibitory outputs from the striatum to the tectum and tegmentum. 7 Visually guided behaviors may be regulated through the inhibitory striato-tegmental pathway found in amphibians in a study performed at the Baylor College of Medicine and the Chinese Academy of Sciences. The basal ganglia in amphibians is very important in receiving visual, auditory, olfactory, and mechansensory inputs; the disinhibitory striato-protecto-tectal pathway is important in prey-catching behaviors of amphibians. When the ipsilateral striatum of an adult toad was electrically stimulated, inhibitory postsynaptic potentials were induced in binocular tegmental neurons, which affects the visual system of the toad.

Studies edit

Inhibitory postsynaptic potentials can be inhibited themselves through a signaling process called depolarized-induced suppression of inhibition DSI in CA1 pyramidal cells and cerebellar Purkinje cells. 8 9 In a laboratory setting step depolarizations the soma have been used to create DSIs, but it can also be achieved through synaptically induced depolarization of the dendrites. DSIs can be blocked by ionotropic receptor calcium ion channel antagonists on the somata and proximal apical dendrites of CA1 pyramidal cells. Dendritic inhibitory postsynaptic potentials can be severely reduced by DSIs through direct depolarization.

Along these lines, inhibitory postsynaptic potentials are useful in the signaling of the olfactory bulb to the olfactory cortex. 10 EPSPs are amplified by persistent sodium ion conductance in external tufted cells. Low-voltage activated calcium ion conductance enhances even larger EPSPs. The hyperpolarization activated nonselective cation conductance decreases EPSP summation and duration and they also change inhibitory inputs into postsynaptic excitation. IPSPs come into the picture when the tufted cells membranes are depolarized and IPSPs then cause inhibition. At resting threshold IPSPs induce action potentials. GABA is responsible for much of the work of the IPSPs in the external tufted cells.

Another interesting study of inhibitory postsynaptic potentials looks at neuronal theta rhythm oscillations that can be used to represent electrophysiological phenomena and various behaviors. 11 12 Theta rhythms are found in the hippocampus and GABAergic synaptic inhibition helps to modulate them. They are dependent on IPSPs and started in either CA3 by muscarinic acetylcholine receptors and within C1 by the activation of group I metabotropic glutamate receptors. When interneurons are activated by metabotropic acetylcholine receptors in the CA1 region of rat hippocampal slices, a theta pattern of IPSPs in pyramidal cells occurs independent of the input. This research also studies DSIs, showing that DSIs interrupt metabotropic acetylcholine-initiated rhythm through the release of endocannabinoids. An endocannabinoid-dependent mechanism can disrupt theta IPSPs through action potentials delivered as a burst pattern or brief train. In addition, the activation of metabotropic glutamate receptors removes any theta IPSP activity through a G-protein, calcium ion–independent pathway.

Inhibitory postsynaptic potentials have also been studied in the Purkinje cell through dendritic amplification. The study focused in on the propagation of IPSPs along dendrites and its dependency of ionotropic receptors by measuring the amplitude and time-course of the inhibitory postsynaptic potential. The results showed that both compound and unitary inhibitory postsynaptic potentials are amplified by dendritic calcium ion channels. The width of a somatic IPSP is independent of the distance between the soma and the synapse whereas the rise time increases with this distance. These IPSPs also regulate theta rhythms in pyramidal cells. On the other hand, inhibitory postsynaptic potentials are depolarizing and sometimes excitatory in immature mammalian spinal neurons because of high concentrations of intracellular chloride through ionotropic GABA or glycine chloride ion channels. 13 These depolarizations activate voltage-dependent calcium channels. They later become hyperpolarizing as the mammal matures. To be specific, in rats, this maturation occurs during the perinatal period when brain stem projects reach the lumbar enlargement. Descending modulatory inputs are necessary for the developmental shift from depolarizing to hyperpolarizing inhibitory postsynaptic potentials. This was studied through complete spinal cord transections at birth of rats and recording IPSPs from lumbar motoneurons at the end of the first week after birth.

Glutamate, an excitatory neurotransmitter, is usually associated with excitatory postsynaptic potentials in synaptic transmission. However, a study completed at the Vollum Institute at the Oregon Health Sciences University demonstrates that glutamate can also be used to induce inhibitory postsynaptic potentials in neurons. 14 This study explains that metabotropic glutamate receptors feature activated G proteins in dopamine neurons that induce phosphoinositide hydrolysis. The resultant products bind to inositol triphosphate IP3 receptors through calcium ion channels. The calcium comes from stores and activate potassium conductance, which causes a pure inhibition in the dopamine cells. The changing levels of synaptically released glutamate creates an excitation through the activation of ionotropic receptors, followed by the inhibition of metabotropic glutamate receptors.