Sleeper Cell: Preliminary Chapter (Sleeper Cell 2240)
The nursing facility shall also provide access to and a copy of other information from the resident's record that has been requested by or on behalf of the resident to support a resident's reconsideration request. Notwithstanding any law to the contrary, the facility may not charge a fee for providing copies of the requested documentation. This notice must inform the resident or the resident's representative that a reconsideration of the resident's classification is being requested, the reason for the request, that the resident's rate will change if the request is approved by the commissioner, the extent of the change, that copies of the facility's request and supporting documentation are available for review, and that the resident also has the right to request a reconsideration.
If the facility fails to provide the required information listed in item iii with the reconsideration request, the commissioner may request that the facility provide the information within 14 calendar days. The reconsideration request must be denied if the information is then not provided, and the facility may not make further reconsideration requests on that specific reimbursement classification. The reconsideration must be based upon the assessment that determined the classification and upon the information provided to the commissioner under paragraphs a and b. The original classification must be modified if the commissioner determines that the assessment resulting in the classification did not accurately reflect characteristics of the resident at the time of the assessment.
The resident and the nursing facility or boarding care home shall be notified within five working days after the decision is made. A decision by the commissioner under this subdivision is the final administrative decision of the agency for the party requesting reconsideration.
If a request for reconsideration applies to an assessment used to determine nursing facility level of care under subdivision 4, paragraph c , the resident shall continue to be eligible for nursing facility level of care while the request for reconsideration is pending. The commissioner shall reclassify a resident if the commissioner determines that the resident was incorrectly classified.
The commissioner may also observe and speak to facility staff and residents. If a facility has two successive audits in which the percentage of change is five percent or less and the facility has not been the subject of a special audit in the past 36 months, the facility may be audited biannually. A stratified sample of 15 percent, with a minimum of ten assessments, of the most current assessments shall be selected for audit.
If more than 20 percent of the RUG-IV classifications are changed as a result of the audit, the audit shall be expanded to a second 15 percent sample, with a minimum of ten assessments. If the total change between the first and second samples is 35 percent or greater, the commissioner may expand the audit to all of the remaining assessments. If a facility has two expanded audits within a month period, that facility will be audited at least every six months for the next 18 months.
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These circumstances include, but are not limited to, the following:. If the results of the audit reflect a change in the resident's case mix classification, a case mix classification notice will be made available electronically to the facility, using the procedure in subdivision 7, paragraph a. The notice must contain the resident's classification and a statement informing the resident, the resident's authorized representative, and the facility of their right to review the commissioner's documents supporting the classification and to request a reconsideration of the classification.
This notice must also include the address and telephone number of the Office of Ombudsman for Long-Term Care. After implementation of this section, reconsiderations requested for classifications made under section The person is considered at risk under this clause if the person currently lives alone or will live alone or be homeless without the person's current housing and also meets one of the following criteria:.
In no case shall medical assistance payment for long-term care services occur prior to the date of the determination of nursing facility level of care. The notice shall include the following information:. A recipient who meets the criteria in section B. This paragraph is in effect for appeals filed between January 1, , and December 31, Subdivision 11 is effective on or after January 1, , for individuals age 21 and older, and on or after October 1, , for individuals under age Laws , chapter 79, article 8, section 4, the effective date, as amended by Laws , First Special Session chapter 1, article 24, section 12, and Laws , First Special Session chapter 9, article 7, section The state commissioner of health may receive and accept money, property, or services from any person, agency, or other source for any public health purpose within the scope of statutory authority.
All money so received is annually appropriated for those purposes in the manner and subject to the provisions of law applicable to appropriations of state funds. The commissioner of health is authorized to enter into contractual agreements with any public or private entity for the provision of statutorily prescribed public health services by the department. The contracts shall specify the services to be provided and the amount and method of reimbursement therefor.
Funds generated in a contractual agreement made pursuant to this section are appropriated to the department for purposes of providing the services specified in the contracts. All such contractual agreements shall be processed in accordance with the provisions of chapter 16C. All mobile health evaluation and screening providers must be directly supervised by a physician licensed under chapter A mobile health evaluation and screening provider shall be required to comply with all licensing reporting and certification, sanitation, and other requirements and regulations that apply to a health care provider supplying similar services as a fixed location provider.
A mobile health evaluation and screening provider shall be subject to regulation and order of the Department of Health. A mobile health evaluation and screening provider shall register with the commissioner and file the anticipated locations of practice, schedules, and routes annually no later than January The mobile health evaluation and screening provider shall also include the name and address of the supervising physician.
A mobile health evaluation and screening provider shall provide at least 30 days' written notice to the populations they intend to serve. The commissioner of health may develop and coordinate a reporting system to improve the state's ability to document inadequate nutrient and food intake of Minnesota's children and adults and to identify problems and determine the most appropriate strategies for improving inadequate nutritional status.
The Board on Aging may develop a method to evaluate the nutritional status and requirements of the elderly in Minnesota. The commissioner of management and budget is hereby appointed as the custodian of all moneys received, or which may hereafter be received, by the state by reason of any federal aid granted for maternal and child welfare service and for public health services, including the purposes as declared in Public Law enacted by the 79th Congress of the United States, Chapter d Session and all amendments thereto, which moneys shall be expended in accordance with the purposes expressed in the acts of Congress granting such aid and solely in accordance with plans to be prepared by the state commissioner of health.
The plans so to be prepared by the commissioner of health for maternal and child health service shall be approved by the United States Children's Bureau; and the plans of the commissioner of health for public health service shall be approved by the United States Public Health Service.
Such plans shall include the training of personnel for both state and local health work and conform with all the requirements governing federal aid for these purposes. Such plans shall be designed to secure for the state the maximum amount of federal aid which is possible to be secured on the basis of the available state, county, and local appropriations for such purposes.
The commissioner of health shall make reports, which shall be in such form and contain such information as may be required by the United States Children's Bureau or the United States Public Health Service, as the case may be; and comply with all the provisions, rules, and regulations which may be prescribed by these federal authorities in order to secure the correction and verification of such reports.
The commissioner may make such reasonable rules as may be necessary to carry into effect the provisions of section Three weeks' published notice of such rules, if of general application throughout the state, shall be given at the seat of government; if of local application only, as near such locality as practicable.
Special rules applicable to particular cases shall be sufficiently noticed when posted in a conspicuous place upon or near the premises affected. Fines collected for violations of rules adopted by the commissioner shall be paid into the state treasury; and of local boards and officers, into the county treasury. The commissioner may adopt reasonable rules pursuant to chapter 14 for the preservation of the public health. The rules shall not conflict with the charter or ordinance of a city of the first class upon the same subject.
The commissioner may control, by rule, by requiring the taking out of licenses or permits, or by other appropriate means, any of the following matters:. Neither the commissioner nor any community health board as defined in section A. These laboratories shall not include laboratories owned or operated by five or less licensed practitioners of the healing arts, unless otherwise provided by federal law or regulation, and in which these practitioners perform tests or procedures solely in connection with the treatment of their patients. Rules promulgated under the authority of this clause, which shall not take effect until federal legislation relating to the regulation and improvement of clinical laboratories has been enacted, may relate at least to minimum requirements for external and internal quality control, equipment, facility environment, personnel, administration and records.
These rules may include the establishment of a fee schedule for clinical laboratory inspections. The provisions of this clause shall expire 30 days after the conclusion of any fiscal year in which the federal government pays for less than 45 percent of the cost of regulating clinical laboratories. The commissioner may regulate the general sanitation of mass gatherings by promulgation of rules in respect to, but not limited to, the following areas: No permit, license, or other prior approval shall be required of the commissioner for a mass gathering.
A "mass gathering" shall mean an actual or reasonably anticipated assembly of more than 1, persons which will continue, or may reasonably be expected to continue, for a period of more than ten consecutive hours and which is held in an open space or temporary structure especially constructed, erected or assembled for the gathering. For purposes of this subdivision, "mass gatherings" shall not include public gatherings sponsored by a political subdivision or a nonprofit organization. Applications for licenses or permits issued pursuant to this section shall be submitted with a fee prescribed by the commissioner pursuant to section Licenses or permits shall expire and be renewed as prescribed by the commissioner pursuant to section Camper cabins and bunkhouses are exempt from floor space, air space, or bed spacing requirements applicable to lodging establishments adopted by the commissioner.
For the purposes of this section:. For purposes of sections Radioactive material includes special nuclear material, source nuclear material, and by-product nuclear material. Source nuclear material does not include special nuclear material. In order to have a comprehensive program to protect the public from radiation hazards, the governor, on behalf of the state, is authorized to enter into agreements with the United States Nuclear Regulatory Commission under the Atomic Energy Act of , section b, as amended.
The agreement shall provide for the discontinuance of portions of the Nuclear Regulatory Commission's licensing and related regulatory authority over by-product, source, and special nuclear materials, and the assumption of regulatory authority over these materials by the state. The Department of Health is designated as the lead agency to pursue an agreement on behalf of the governor and for any assumption of specified licensing and regulatory authority from the Nuclear Regulatory Commission under an agreement with the commission.
The commissioner of health shall establish an advisory group to assist in preparing the state to meet the requirements for reaching an agreement.
Neurophysiological and Computational Principles of Cortical Rhythms in Cognition
The commissioner may adopt rules to allow the state to assume regulatory authority under an agreement under this section, including the licensing and regulation of radioactive materials. Any regulatory authority assumed by the state includes the ability to set and collect fees. A person who, on the effective date of an agreement under this section, possesses a Nuclear Regulatory Commission license that is subject to the agreement is deemed to possess a similar license issued by the Department of Health.
A Department of Health license obtained under this subdivision expires on the expiration date specified in the federal license. The commissioner shall adopt rules to ensure that individuals handling or utilizing radioactive materials under the terms of a license issued by the commissioner under section The rules adopted must be at least as stringent as federal regulations on proper training and qualifications adopted by the Nuclear Regulatory Commission.
Rules adopted under this section may incorporate federal regulations by reference. The commissioner may require an applicant for a license under section The financial assurances posted must be sufficient to restore the site to unrestricted future use and must be sufficient to provide for surveillance and care when radioactive materials remain at the site after the licensed activities cease. The commissioner may establish financial assurance criteria by rule. In establishing such criteria, the commissioner may consider:. The commissioner may, by rule, establish types of financial assurances that meet the requirements of this section.
Such financial assurances may include bank letters of credit, deposits of cash, or deposits of government securities. Financial assurances must be established together with trust agreements. Both the financial assurances and the trust agreements must be in a form and substance that meet requirements established by the commissioner. The commissioner is authorized to exempt from the requirements of this section, by rule, any category of licensee upon a determination by the commissioner that an exemption does not result in a significant risk to the public health or safety or to the environment and does not pose a financial risk to the state.
Nothing in this section relieves a licensee of a civil liability incurred, nor may this section be construed to relieve the licensee of obligations to prevent or mitigate the consequences of improper handling or abandonment of radioactive materials. When a license is required for radioactive material or source or special nuclear material by a rule adopted under section The licensee must renew the license 60 days before the expiration date of the license.
The expiration date of a license is the date specified by rule of the commissioner of health. A licensee must pay an annual fee at least 60 days before the anniversary date of the issuance of the license. The annual fee is as follows:. Radioactive materials are limited to ten radionuclides not to exceed a total activity amount of one curie. An annual fee or a license renewal fee submitted to the commissioner after the due date specified by rule must be accompanied by an additional amount equal to 25 percent of the fee due.
The commissioner of health shall make periodic safety inspections of the radioactive material and source and special nuclear material of a licensee. The commissioner shall prescribe the frequency of safety inspections by rule. If the commissioner finds serious violations of public health standards during an inspection under subdivision 6, the licensee must pay all costs associated with subsequent reinspection of the source.
The costs shall be the actual costs incurred by the commissioner and include, but are not limited to, labor, transportation, per diem, materials, legal fees, testing, and monitoring costs. For a period of days or more, the licensee must obtain a license under subdivision 4. The fee for the registration for x-ray machines and other sources of ionizing radiation required to be registered under rules adopted by the state commissioner of health pursuant to section The registration shall expire and be renewed as prescribed by the commissioner pursuant to section Applications for initial or renewal registrations submitted to the commissioner after the time specified by the commissioner shall be accompanied by an amount equal to 25 percent of the fee due in addition to the fees prescribed in subdivision 1a.
Periodic radiation safety inspections of the sources of ionizing radiation shall be made by the state commissioner of health. The frequency of safety inspections shall be prescribed by the commissioner on the basis of the frequency of use of the source of ionizing radiation; provided that each source shall be inspected at least once every four years. Notwithstanding rules adopted by the commissioner under section Equivalent examinations may be approved by the commissioner, if the examination is consistent with the standards for educational and psychological testing as recommended by the American Education Research Association, the American Psychological Association, the National Council on Measurement in Education, or the National Commission for Certifying Agencies.
The collected fee shall be deposited in the state treasury and credited to the state government special revenue fund. The examinations must include the anatomy of, and positioning for, the specific regions. The processing fee and the examination fee shall be deposited in the state treasury and credited to the state government special revenue fund.
The commissioner shall submit the fee to the national organization providing the examination. The operator may practice using only routine radiographic procedures, for the interpretation by and under the direction of a licensed practitioner, excluding computed tomography, the use of contrast media, and the use of fluoroscopic or mammographic equipment. The commissioner may grant a variance according to Minnesota Rules, parts The request for a variance must be in writing, state the circumstances that constitute hardship, state the period of time the facility wishes to have the variance for the scope of practice in place, and state the alternative measures that will be taken if the variance is granted.
The commissioner shall set forth in writing the reasons for granting or denying the variance. Variances granted by the commissioner must specify in writing the time limitation and required alternative measures to be taken by the facility. A request for the variance shall be denied if the commissioner finds the circumstances stated by the facility do not support a claim of hardship, the requested time period for the variance is unreasonable, the alternative measures proposed by the facility are not equivalent to the scope of practice, or the request for the variance is not submitted to the commissioner in a timely manner.
At the time a facility with x-ray equipment is inspected by the commissioner of health in accordance with subdivision 2, an individual operating x-ray equipment in the facility must be able to show compliance with the requirements of subdivision 5. For purposes of this section, "facility" has the meaning provided in United States Code, title 42, section b a 3 A. A facility at which a mammography examination is performed shall, if a patient is categorized by the facility as having heterogeneously dense breasts or extremely dense breasts based on the Breast Imaging Reporting and Data System established by the American College of Radiology, include in the summary of the written report that is sent to the patient, as required by the federal Mammography Quality Standards Act, United States Code, title 42, section b, notice that the patient has dense breast tissue, that this may make it more difficult to detect cancer on a mammogram, and that it may increase her risk of breast cancer.
The following language may be used:. Dense breast tissue is relatively common and is found in more than 40 percent of women. However, dense breast tissue may make it more difficult to identify precancerous lesions or cancer through a mammogram and may also be associated with an increased risk of breast cancer.
This information about the results of your mammogram is given to you to raise your own awareness and to help inform your conversations with your treating clinician who has received a report of your mammogram results. Together you can decide which screening options are right for you based on your mammogram results, individual risk factors, or physical examination. For purposes of this section, "handheld dental x-ray equipment" means x-ray equipment that is used to take dental radiographs, is designed to be handheld during operation, and is operated by an individual authorized to take dental radiographs under chapter A.
Calibration must include the test specified in Minnesota Rules, part Handheld dental x-ray equipment used according to this section and according to manufacturer instructions is exempt from the following requirements for the equipment:. A registrant using handheld dental x-ray equipment shall otherwise comply with Minnesota Rules, chapter The expiration dates of the various licenses, permits, registrations, and certifications as prescribed by the rules shall be plainly marked thereon.
Fees may include application and examination fees and a penalty fee for renewal applications submitted after the expiration date of the previously issued permit, license, registration, and certification. The commissioner may also prescribe, by rule, reduced fees for permits, licenses, registrations, and certifications when the application therefor is submitted during the last three months of the permit, license, registration, or certification period.
Fees proposed to be prescribed in the rules shall be first approved by the Department of Management and Budget. All fees proposed to be prescribed in rules shall be reasonable. The fees shall be in an amount so that the total fees collected by the commissioner will, where practical, approximate the cost to the commissioner in administering the program. All fees collected shall be deposited in the state treasury and credited to the state government special revenue fund unless otherwise specifically appropriated by law for specific purposes.
Fees charged for environment and medical laboratory services provided by the department must be approximately equal to the costs of providing the services. All receipts generated by the program are annually appropriated to the commissioner for use in the maternal and child health program. The commissioner shall set license fees for outpatient surgical centers, boarding care homes, and supervised living facilities at the following levels:. Fees collected under this paragraph are nonrefundable. The fees are nonrefundable even if received before July 1, , for licenses or registrations being issued effective July 1, , or later.
These fees shall be submitted at the time of the application for federal certification and shall not be refunded. All fees collected after the date that the imposition of fees is not prohibited by federal law shall be deposited in the state treasury and credited to the state government special revenue fund. The commissioner of health shall be responsible for the adoption of rules and enforcement of applicable laws and rules relating to the operation, maintenance, design, installation, and construction of public pools and facilities related to them.
The commissioner shall adopt rules governing the collection of fees under section All plans and specifications for public pool and spa construction, installation, or alteration or requests for a variance that are submitted to the commissioner according to Minnesota Rules, part All public pool construction plans submitted for review after January 1, , must be certified by a professional engineer registered in the state of Minnesota. If the commissioner determines, upon review of the plans, that inadequate fees were paid, the necessary additional fees shall be paid before plan approval.
For purposes of determining fees, a project is defined as a proposal to construct or install a public pool, spa, special purpose pool, or wading pool and all associated water treatment equipment and drains, gutters, decks, water recreation features, spray pads, and those design and safety features that are within five feet of any pool or spa.
The commissioner shall charge the following fees for plan review and inspection of public pools and spas and for requests for variance from the public pool and spa rules:. The record required under Minnesota Rules, part The pool must not be reopened until the missing or broken cover or grate has been replaced according to the manufacturer's specifications, or the loose cover or grate has been reattached to the manufacturer's specifications. A pool used by a medical or rehabilitation facility to facilitate treatment or therapy, to which only authorized access is allowed and which is not open for any other public use, is exempt from the requirements of Minnesota Rules, part A portable wading pool that is located at a family day care or group family day care home licensed under Minnesota Rules, chapter , or at a home at which child care services are provided under section A.
If the provider chooses to allow children cared for at the family day care or group family day care home to use the swimming pool located at the home, the provider must satisfy the requirements in section A. This notice is required under Minnesota Statutes, section The commissioner of health shall be responsible for the adoption of rules and enforcement of applicable laws and rules relating to indoor air quality in the operation and maintenance of enclosed sports arenas.
For purposes of this section, "advanced diagnostic imaging services" has the meaning given in United States Code, title 42, section M, except that it does not include x-ray, ultrasound, or fluoroscopy. Thereafter, all facilities that provide advanced diagnostic imaging services in the state must obtain licensure or accreditation within six months of commencing operations and must maintain either licensure pursuant to sections Except for the limitation contained in this section, the commissioner of health may enter into a contractual agreement to recover costs incurred for analysis for diagnostic purposes for each specimen submitted to the Department of Health by any hospital, laboratory, clinic, or physician.
The commissioner shall not charge for any biological materials submitted to the Department of Health as a requirement of Minnesota Rules, part The commissioner of health may establish other exceptions to the handling fee as may be necessary to protect the public's health.
Funds generated in a contractual agreement made pursuant to this section shall be deposited in a special account and are appropriated to the commissioner for purposes of providing the services specified in the contracts. This fee amount shall be deposited in the state treasury and credited to the state government special revenue fund.
This fee shall be deposited in the state treasury and credited to the general fund. The commissioner shall periodically revise the list of tests to be administered for determining the presence of a heritable or congenital disorder. Revisions to the list shall reflect advances in medical science, new and improved testing methods, or other factors that will improve the public health.
In determining whether a test must be administered, the commissioner shall take into consideration the adequacy of analytical methods to detect the heritable or congenital disorder, the ability to treat or prevent medical conditions caused by the heritable or congenital disorder, and the severity of the medical conditions caused by the heritable or congenital disorder. The list of tests to be performed may be revised if the changes are recommended by the advisory committee established under section The revision is exempt from the rulemaking requirements in chapter 14, and sections The department shall promote the materials describing the newborn screening program and encourage providers and education programs to thoroughly discuss the program with expectant parents and parents with newborns.
The department shall make information and forms about newborn screening available to the persons with a duty to perform testing under this section and to expectant parents and parents of newborns using electronic and other means. The signed form must be made part of the infant's medical record and a copy shall be provided to the Department of Health. When a parent or legal guardian elects not to have newborn screening performed, the person with the duty to perform testing under subdivision 1 must follow that election. A written election to decline testing exempts persons with a duty to perform testing and the Department of Health from the requirements of this section and section The consent form must provide the following:.
If written informed consent of a parent, legal guardian, or individual is obtained under subdivision 7, the Department of Health may use the blood samples and test results in accordance with subdivision 9. Until destroyed, the blood samples and test results may be used for program operations described under subdivision 5, paragraph a , clauses 1 to 7.
With the written, informed consent of a parent or legal guardian, the Department of Health may use blood samples and test results for public health studies or research not related to newborn screening, and upon approval by the Department of Health's Institutional Review Board, share samples and test results with external parties for public health studies or research. A parent or legal guardian, or the individual whose blood was tested as an infant if the individual is 18 years of age or older, may revoke approval for storage or use of blood samples or test results at any time by providing a signed and dated form requesting destruction of the blood samples or test results.
Blood samples and test results must be destroyed as specified under subdivision 8, paragraph b. The screening must occur after the infant is 24 hours old, before discharge from the nursery. If discharge occurs before the infant is 24 hours old, the screening must occur as close as possible to the time of discharge. The information and forms must include screening protocol and reporting requirements and parental options;. Membership of the committee shall include, but not be limited to, at least one member from each of the following representative groups:.
Members shall not receive per diems but shall be compensated for expenses. If the defendant does not move for dissolution of the writ as provided in s. If the property is in another jurisdiction, the party who had the writ issued shall deliver the writ to the sheriff in the other jurisdiction; and that sheriff shall execute the writ, upon order of the court, by levying on the property and delivering it to the sheriff of the county in which the action is pending, to be disposed of according to law, unless he or she is ordered by the court from which the writ emanated to hold the property and dispose of it in his or her jurisdiction according to law.
If the plaintiff shows by a sworn statement that the defendant cannot be found within the state, the levy on the property suffices as service on the defendant. The court shall hear the motion not later than the day on which the sheriff is authorized under the writ to levy on property liable under distress. If the plaintiff proves a prima facie case, or if the defendant defaults, the court shall order the sheriff to proceed with the levy.
The bond shall be approved by such sheriff; made payable to plaintiff in double the value of the property levied on, with the value to be fixed by the sheriff; and conditioned for the forthcoming of the property restored to abide the final order of the court. Judgment may be entered against the surety on such bonds in the manner and with like effect as provided in s. If the verdict or the finding of the court is for defendant, the action shall be dismissed and defendant shall have judgment and execution against plaintiff for costs.
The service of the notice shall be by delivery of a true copy thereof, or, if the tenant is absent from the rented premises, by leaving a copy thereof at such place. This subsection applies only when the lease is silent on the matter or when the tenancy is an oral one at will. The notice may give a longer time period for cure of the breach or surrender of the premises. In the absence of a lease provision prescribing the method for serving notices, service must be by mail, hand delivery, or, if the tenant is absent from the rental premises or the address designated by the lease, by posting.
The tenant shall serve the landlord, in the manner prescribed by s. The lease may provide for a longer period of time for repair or maintenance. Once the landlord has completed the repair or maintenance, the tenant shall pay the landlord the amounts of rent withheld. If the landlord does not complete the repair or maintenance in the allotted time, the parties may extend the time by written agreement or the tenant may abandon the premises, retain the amounts of rent withheld, terminate the lease, and avoid any liability for future rent or charges under the lease.
This section is cumulative to other existing remedies, and this section does not prevent any tenant from exercising his or her other remedies.
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Acceptance of the rent includes conduct by the landlord concerning any tender of the rent by the tenant which is inconsistent with reasonably prompt return of the payment to the tenant. The minimum time delay between the two attempts to obtain service shall be 6 hours. One envelope shall be addressed to such address or location as has been designated by the tenant for receipt of notice in a written lease or other agreement or, if none has been designated, to the residence of the tenant, if known. The second envelope shall be addressed to the last known business address of the tenant.
The clerk of the court shall immediately mail the copies of the summons and complaint by first-class mail, note the fact of mailing in the docket, and file a certificate in the court file of the fact and date of mailing. Service shall be effective on the date of posting or mailing, whichever occurs later; and at least 5 days from the date of service must have elapsed before a judgment for final removal of the defendant may be entered. If the plaintiff expressly and specifically sought money damages in the complaint, in addition to awarding possession of the premises to the plaintiff, the court shall also direct, in an amount which is within its jurisdictional limitations, the entry of a money judgment in favor of the plaintiff and against the defendant for the amount of money found due, owing, and unpaid by the defendant, with costs.
However, no money judgment shall be entered unless service of process has been effected by personal service or, where authorized by law, by certified or registered mail, return receipt, or in any other manner prescribed by law or the rules of the court, and no money judgment may be entered except in compliance with the Florida Rules of Civil Procedure. If the issues are found for defendant, judgment shall be entered dismissing the action. Unless the tenant disputes the amount of accrued rent, the tenant must pay the amount alleged in the complaint into the court registry on or before the date on which his or her answer to the claim for possession is due.
If the tenant contests the amount of accrued rent, the tenant must pay the amount determined by the court into the court registry on the day that the court makes its determination. The court may, however, extend these time periods to allow for later payment, upon good cause shown. Even though the defense of payment or satisfaction has been asserted, the court, in its discretion, may order the tenant to pay into the court registry the rent that accrues during the pendency of the action, the time of accrual being as set forth in the lease.
If the landlord is in actual danger of loss of the premises or other hardship resulting from the loss of rental income from the premises, the landlord may apply to the court for disbursement of all or part of the funds so held in the court registry. In such case, the landlord is entitled to an immediate default for possession without further notice or hearing thereon. For residents of a facility licensed under part II of chapter , the provisions of s.
An early termination fee does not include: If the rent is payable weekly, then the tenancy is from week to week; if payable monthly, tenancy is from month to month; if payable quarterly, tenancy is from quarter to quarter; if payable yearly, tenancy is from year to year. If wages are payable weekly or more frequently, then the tenancy is from week to week; and if wages are payable monthly or no wages are payable, then the tenancy is from month to month.
In the event that the employee ceases employment, the employer shall be entitled to rent for the period from the day after the employee ceases employment until the day that the dwelling unit is vacated at a rate equivalent to the rate charged for similarly situated residences in the area. This subsection shall not apply to an employee or a resident manager of an apartment house or an apartment complex when there is a written agreement to the contrary.
The right to attorney fees in this section may not be waived in a lease agreement. However, attorney fees may not be awarded under this section in a claim for personal injury damages based on a breach of duty under s. The landlord shall not commingle such moneys with any other funds of the landlord or hypothecate, pledge, or in any other way make use of such moneys until such moneys are actually due the landlord;. The landlord shall not commingle such moneys with any other funds of the landlord or hypothecate, pledge, or in any other way make use of such moneys until such moneys are actually due the landlord; or.
In addition to posting the surety bond, the landlord shall pay to the tenant interest at the rate of 5 percent per year, simple interest. In addition to posting a surety bond, the landlord shall pay to the tenant interest on the security deposit or advance rent held on behalf of that tenant at the rate of 5 percent per year simple interest. Subsequent to providing such written notice, if the landlord changes the manner or location in which he or she is holding the advance rent or security deposit, he or she must notify the tenant within 30 days after the change as provided in paragraphs a - d.
The landlord is not required to give new or additional notice solely because the depository has merged with another financial institution, changed its name, or transferred ownership to a different financial institution. This subsection does not apply to any landlord who rents fewer than five individual dwelling units.
Failure to give this notice is not a defense to the payment of rent when due. The written notice must: For all other deposits: The notice shall contain a statement in substantially the following form: The failure of the tenant to make a timely objection does not waive any rights of the tenant to seek damages in a separate action. The court shall advance the cause on the calendar. Enforcement personnel shall look solely to this section to determine compliance. This section prevails over any conflicting provisions in chapter and in other sections of the Florida Statutes, and shall operate to permit licensed real estate brokers to disburse security deposits and deposit money without having to comply with the notice and settlement procedures contained in s.
With the exception of subsections 3 , 5 , and 6 , this section is not applicable to housing authorities or public housing agencies created pursuant to chapter or other statutes. Failure to give such notice shall relieve the landlord of the notice requirement of paragraph 3 a but shall not waive any right the tenant may have to the security deposit or any part of it. The parameter changes reduce the potassium conductance and speed up the sodium and potassium current kinetics.
As a result, the recovery from hyperpolarization is fast, and the delay portion of the PRC essentially disappears. Therefore, type I and type II PRCs can be produced by the same set of ion channels but with quantitatively different properties. For type I neurons, a cell that fires earlier is phase-advanced by another cell that fires later, which acts against reducing the relative phase difference between the two.
Hence it is not straightforward to ascertain intuitively synchronization by mutual excitation to type I neurons. As it turns out, a mathematical analysis using the phase reduction theory, which will be discussed in Section III, found that type I neurons tend to be much more easily synchronized by mutual inhibition than excitation [ , , ].
This theoretical finding shows that single cell properties can qualitatively alter network behavior in a completely counter-intuitive way. Phase response properties have been routinely characterized for neurons in central pattern generator systems [ , ]. More recently, in vitro slice studies have directly assessed PRCs of neurons in the mammalian cerebral cortex, using current perturbations that mimic excitatory or inhibitory postsynaptic potential EPSP or IPSP [ , , , ]. It was generally found that for a given cell, the classification of its PRC into type I or type II is consistent for a range of intrinsic oscillation frequencies, and not sensitive to the exact amplitude of PSP perturbations [ , ].
Distinct neuron types exhibit differential preponderance for phase response properties. Nevertheless, neurons within a neural population are heterogeneous and exhibit both type I and type II PRCs [ ], possibly as a result of quantitative differences in the expression of ion channels across neurons in a population. The precise shape of PRC of a neuron depends on its repertoire and somatodendritic distribution of various ion channels. In general, regenerative currents such as a persistent sodium current I NaP that provides a positive feedback for membrane depolarization tend to shift leftward a PRC and eradicate the negative lobe, thereby favoring type I behavior [ , ].
On the other hand, restorative currents such as a voltage-activated M-type potassium current I M that acts as a negative feedback for voltage changes tend to accentuate the negative lobe [ , , , ]. This is because a current like I M is activated by depolarization during an action potential, and decays afterwards as the membrane recovers from afterhyperpolarization.
A brief depolarization soon after a spike would counter this decay process, hence delaying the timing of the next spike. The hyperpolarization activated cation current I H de-activates with depolarization, hence has a similar effect as a depolarization-activated outward current. I H is thus expected to accentuate the negative lobe and favors type II PRC behavior, a prediction that was confirmed with pharmacologically blockade of I H in pyramidal neurons [ ].
Another important issue is how PRC depends on the location of synaptic inputs in a spatially complex neuron [ , ]. As a result, somatic and dendritic PRCs can in principle show disparate behaviors. For example, because of an increasing gradient of expression of the H ion channel conductance from soma to distal dendrite [ ], dendrites of a pyramidal cell can display type II PRC, whereas its somatic PRC can be type I. A calcium-activated potassium current I KCa and, to a lesser degree, I M and I H , underlie spike-frequency adaptation, which is a common characteristic of pyramidal neurons [ , , , , , , ] and certain subclasses of inhibitory cells [ , ].
As it turns out, a large I M which has a low voltage threshold and is significant at resting membrane potential , but not I KCa which depends on calcium influx through the opening of high-threshold calcium channels during action potentials , can switch a neuron from type I to type II [ , ].
Conversely, a reduction of I M e. Indeed, in vitro pyramidal cells display strong spike-frequency adaptation, yet show mostly type I behavior; when they do display type II PRCs the negative lobe is fairly shallow [ , , , ]. Such differential PRC properties of different neural types have important implications for their distinct roles in generating network rhythms see Section III. Resonance refers to the phenomenon of band-passed neuronal response, with maximal response at a preferred input frequency. Resonance is intuitively expected when single neurons are damped oscillators hence display an intrinsic preferred frequency [ , ].
However, resonance does not necessarily require damped oscillation of single neurons [ , ]. Membrane passive properties e. For example, an I H opposes voltage changes in a subthreshold voltage range: I H has a time constant on the order of a hundred milliseconds, thus behaves as a high-pass filter. An H-current dependent mechanism underlies resonance observed in layer II stellate cells of entorhinal cortex, and combined with membrane noise gives rise to subthreshold oscillations in these cells [ , , , , , ].
Interestingly, I H -dependent resonance is most prominent when the inputs are located at distal dendrites [ , , , ], consistent with an increasing gradient of I H expression from soma to distal dendrite of pyramidal neurons [ ]. Another example is a voltage-gated potassium current, like I M or a low-threshold slowly inactivating potassium current I KS , which counter-acts membrane potential changes in a depolarized range.
Together with suppressed responses at high frequencies by the leak conductance, such a current produces resonance when neurons are excited to significantly above the resting membrane potential but below firing threshold [ , ]. A single neuron may display more than one resonance bandpass filters that are spatially segregated, for instance an I H dependent mechanism in the dendrite and an I M dependent mechanism in the soma [ ]. How does subthreshold resonance relate to neuronal firing of action potentials?
It has been shown that, in hippocampus, neurons display similar frequency preference for membrane potential responses and spike discharges at low firing rates when V m stays below firing threshold most of the time [ ]. Moreover, resonance frequency is in the theta frequency range for pyramidal cells, and in the gamma frequency range for fast spiking interneurons, suggesting differential roles of these two major cell types in theta- versus gamma-frequency network oscillations [ ]. In a systematical modeling study of neuronal resonance, Richardson et al [ ] considered two model neurons with resonance frequency at f R , one based on I H and the other on I KS and I NaP.
They used input currents composed of three components: They found that, interestingly, subthreshold resonance may or may not be reflected in the spike firing activity, depending on the degree of neural firing variability. This is true even when the mean firing rate r 0 is much higher than f R , e.
Although somewhat surprising, this result may be explained by observing that, with a stronger noise, a lower mean input is needed to produce the same mean firing rate r 0 , because spikes are more often triggered by stochastic input fluctuations. Therefore on average the membrane potential is lower where subthreshold resonance plays a more important role.
In other words, noise is important for unmasking a subthreshold resonance in the spike firing patterns. This insight is likely to have implications for understanding the relationship between highly irregular neural discharges and coherent network rhythms Section V. A Response with low noise. B Response with high noise. Same format as in A. Phase shift of the firing response relative to the sinusoidal input positive: Adapted with permission from [ ].
A neuron becomes a pacemaker if it displays sustained intrinsic oscillations [ , 15 , , , ]. Oscillation naturally occurs as a result of the interaction between a positive feedback between V m depolarization and I NaP activation and a slower negative feedback depolarization induced I KS activation. The activation time constant of I KS controls the recovery phase of the oscillation cycle hence largely determines the rhythmic frequency which can be in the gamma or theta frequency range [ ].
The same model was analyzed in the study of resonance by Richardson et al. Notably, the model of [ ] predicts a different relationship between the oscillation frequency and the input drive in these two cases. In the gamma frequency regime, subthreshold oscillation frequency increases with the input drive see Section II-C. In the theta frequency regime, the frequency remains roughly constant exhibiting a plateau as long as the oscillation is subthreshold, and starts to increase with the applied current intensity when spike firing becomes significant.
The predicted plateau phenomenon was observed in stellate cells of entorhinal cortex Fig. Among neural types that exhibit subthreshold membrane oscillations, of special interest are GABAergic neurons in the nucleus basalis [ 16 ] and medial septum [ ] Fig. The nucleus basalis and medial septum are the major parts of the basal forebrain that provide the neuromodulatory transmitter acetylcholine to the neocortex and hippocampus, respectively [ , , , ].
In addition to cholinergic cells, these structures also contain GABAergic cells with axonal projections to the cortical mantle. GABAergic cells in the medial septum are believed to act as a pacemaker for theta rhythm in the hippocampus, and those in the nucleus basalis may play a similar role for theta rhythm in the neocortex. As shown in Fig. The inter-cluster oscillation is in the theta frequency range, whereas both subthreshold membrane potential oscillations and intra-cluster spike firing rate are in the gamma frequency range.
Such mixed-mode oscillations are especially interesting because in the entorhinal cortex and hippocampus theta rhythm is often temporally nested with faster gamma-frequency oscillations [ , , , , , , , , ]. A biophysical basis of such inter-nested gamma and theta mixed-mode oscillation was described by the aforementioned ionic channel model [ , ] Fig. The mechanism relies on two assumptions. Second, the neuronal spike afterhyperpolarization is strong, so that the average membrane potential is lower during spike discharges than during subthreshold epochs.
Thus, when the neuron fires a cluster of spikes, the de-inactivation of I KS slowly builds up during spike afterhyperpolarizations, so the amplitude of I KS increases gradually and eventually becomes large enough to terminate a spiking episode. When the cell does not fire spikes in a subsequent subthreshold epoch, the membrane potential is relatively depolarized and I KS slowly decreases due to inactivation, until the cell is sufficiently recovered and can start to fire again. In this scenario, the theta frequency of intrinsic oscillations in septal GABAergic cells is largely controlled by a single current I KS.
This model prediction has not yet been tested experimentally. I KS gradually de-inactivates increase of q , hence I KS during the hyperpolarizing phases of spikes arrows to the right , whereas it inactivates decreasing q during subthreshold membrane oscillations arrows to the left. C Similar behavior of principal mitral cells of the rat olfactory bulb. A Reproduced with permission from [ ]; B from [ ]; C from [ ]. Interestingly, in recent years such subthreshold oscillations have been observed in many cell types, including neocortical neurons [ ], mitral cells in the olfactory bulb [ , 48 ], magnocellular neuron in hypothalamus [ 90 ], mesencephalic trigeminal neurons [ ], dorsal column nuclei neurons in culture [ ], hippocampal interneurons in the lacunosum-moleculare layer [ ].
Similar to the model Fig. In all these diverse cell types, V m oscillations were shown to be blocked by TTX but not by calcium channel blockers, confirming the model prediction that subthreshold oscillations are independent of calcium channels. In this case, spikes are triggered by noise, on top of the membrane potential oscillatory wave [ ]. Also, whereas some cell types are able to exhibit purely subthreshold oscillations [ 13 , , , ], in other neurons subthreshold membrane oscillations always occur interspersed with spike clusters [ 16 , , , ].
This difference presumably reflects whether the spike threshold is significantly above, or overlaps with, the voltage range for activation of the low-threshold ion channels responsible for generating membrane potential oscillations [ ]. In either case, with increasing amplitude of input current, the number of spikes per cluster increases, and spike firing becomes dominant over subthreshold oscillatory episodes. Taken together, there is a convergence of evidence in support of a general ion channel mechanism for subthreshold rhythmogenesis. This offers a powerful means for generating membrane oscillations at distinct frequencies, by a selective expression of a subset of these currents in distinct cell types.
Like pacemaker neurons in central pattern generators [ ], subtypes of cortical cells fire bursts of spikes brief clusters of spikes rhythmically at preferred frequencies [ ]. Bursting neurons have the potential to serve as pacemakers for synchronous network oscillations [ ], and bursts of spikes may underlie or enhance neuronal resonance [ , ]. In the cortex, two types of rhythmic bursting neurons have been observed. Hippocampal pyramidal cells [ ] and layer 5 neocortical pyramidal neurons [ ] show a preponderance for slow rhythmic bursting at 4—15 Hz.
Interestingly, bursting occurs more readily, when its underlying inward and outward channels are located in the dendrite, separated from the perisomatic region where action potentials are produced [ , , , ]. Furthermore, phasic burst firing in layer 5 pyramidal neurons [ ] and in dopaminergic neurons [ , , ] depends on the voltage-gated NMDA receptors for glutamate mediated synaptic transmission.
Because the gamma cycle is only tens of milliseconds, potassium currents such as I M and I KCa are likely to be too slow to set the periodicity of fast bursting in chattering cells. It has also been proposed that after-depolarization depends on a calcium- activated cation current [ 27 ]. Interestingly, a TTX-sensitive sodium channel in interplay with a Kv3 channel in the dendrites were also found to underlie oscillatory bursting in a sensory neuron in electric fish [ , , , ], suggesting that this mechanism may be versatile in vertebrates.
Moreover, calcium-dependent processes can modulate, rather than generate, chattering behavior. Since arousal and attention are associated with cholinergic activation of the cortical system [ , , , ], this result suggest a cholinergic mechanism for promoting neocortical gamma oscillations in attentive states. A A chattering neuron recorded in vivo from the cat visual cortex shows rhythmic bursting in the gamma frequency range.
B A model chattering neuron endowed with a ping-pong interplay between two electrotonic compartments. A Reproduced with permission from [ ], B from [ ]. Subthreshold oscillations and repetitive bursting may have different implications for synchronization of coupled neurons. On the other hand, bursts may provide a reliable signal for the rhythmicity to be transmitted across probabilistic and unreliable synapses between neurons [ , , ].
A generic cortical circuit consists of two major cell types: It follows that three types of synchronization mechanisms by chemical synapses are conceivable: More recently, electrical synapses by gap junctions have also been proposed to contribute to neural synchrony. Here I will describe each of these mechanisms in turn.
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For any type of synaptic coupling, a general mathematical approach for studying synchrony of neural oscillators is provided by the theory of phase-coupled oscillators [ , , , ]. The idea is that, if synaptic interactions are relatively weak, their effect is the synaptic current from the presynaptic neuron into the postsynaptic neuron, averaged over the oscillation period.
Mathematically, we have [ , , , ]. The stability of a steady state phase locking is determined by the slope of H odd through the steady state. The phase reduction theory thus provides a general framework for predicting the synchronization properties of neurons that behave as oscillators and are coupled by relatively weak synaptic interactions.
For a higher frequency rhythm, the period is shorter, and faster synaptic kinetics are required to produce synchrony. The same qualitative explanation applies to synchronization of slow rhythmic firing of epileptic discharges characterized by large after-hyperpolarizations in a disinhibited pyramidal cell population [ , , ].
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As shown above, a key insight from theoretical analysis is that in order to determine network synchrony, interactions between spiking neurons cannot be treated as instantaneous, quantitative synaptic time courses matter a great deal. Furthermore, the required synaptic kinetics is different for different oscillation frequencies.
For example, in hippocampal slices, activation of cholinergic muscarinic receptors by carbachol can induce theta frequency oscillation that is independent of synaptic inhibition [ ]. This rhythm, however, is quite different from theta oscillations in behaving animals during spatial navigation, as it will be discussed later Section IV-F. This is the case even when single neurons display intrinsic rhythmic bursting, as shown in a model of coupled chattering cells [ 28 ]. Thus, synaptic excitation mediated by ionotropic glutamate receptors, alone, is likely to be insufficient for generating coherent network oscillations at high frequencies.
With slow synapses relative to an oscillation period , inhibition rather than excitation generates neural synchrony [ , , , ]. For type I neurons, even fast excitation does not lead to zero-phase synchrony [ , ]. Indeed, it was found that type I neurons typically show anti-phase behavior with excitatory coupling [ , , ].
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On the other hand, reciprocal inhibition turned out to be an effective mechanism for neuronal synchrony. This is illustrated in Fig. Wang and Buzsaki [ ] found that, in this network model of GABAergic interneurons, spike firing oscillation frequency of individual neurons can be varied in a broad range, e. Therefore, networks of inhibitory interneurons provide a mechanism for coherent brain oscillations, in particularly gamma rhythm. In the experiment, when metabotropic glutamate receptors were activated pharmacologically, hippocampal slices exhibited oscillatory activity patterns.
Oscillatory IPSPs in the 40 Hz frequency range were observed in simultaneously recorded pyramidal cells. Drugs that slowed down the kinetics of GABA A receptor mediated synaptic currents led to lower oscillation frequency, confirming a model prediction and lending support of an interneuronal network mechanism for gamma oscillations [ , , ].
At first sight it seems paradoxical that reciprocal inhibition can bring coupled neurons to fire spikes synchronously, since spiking in one neuron triggers a synaptic current that hyperpolarizes the membrane potentials of target neurons. As depicted in a classic model [ ], consider a pair of neurons endowed with some slow fatigue process and coupled reciprocally by fast synaptic inhibition.
At any moment one neuron 1 is active and effectively suppresses the other neuron 2. Whether an inhibitory neural network displays release or escape type behavior depends on the details of active neural properties and synaptic kinetics [ , ], and a dynamic clamp experiment suggests that the same circuit can be switched from one to the other mode by neuromodulation [ ].
It is worth emphasizing again that different synaptic subtypes are suitable for synchronizing circuit rhythms at different frequency ranges. Of course, synaptic currents cannot be too slow relative to the oscillation cycle, otherwise interactions between neurons would become roughly tonic and phasic information critical for synchronization would be lost [ , ]. For type I neurons, synchrony can be realized even with relatively fast inhibitory synapses [ ].
Synaptic latency including axonal delay [ , ] and rise-time [ , , 33 ], as well as the decay time, contribute to determining the synchronous firing patterns. These theoretical results are supported by the observation that, during development of the rodent brain, the frequency and synchrony of gamma oscillations reach adult levels in a time course that parallels with that of cellular properties of basket cells and kinetic time constants of their synaptic connections [ , ]. Just as it is difficult to synchronize an array of clocks with a wide range of intrinsic periods, neural synchrony is more difficult to achieve in the presence of heterogeneity, which introduces a variety of intrinsic oscillation frequencies among individual neurons [ , ].
This is an important issue because the oscillation frequency of a neuron varies considerably with the input current intensity, with type I neurons typically showing a much wider range of firing frequencies than type II neurons [ , ]. This implies that a small variation in the net input current leads to a large change in the oscillation frequency. In vivo , excitatory synaptic currents evoked by sensory stimulation were estimated to be in the range of 0.
Given such input variations, and the high heterogeneity of both cellular and synaptic properties in interneurons [ ], how can synchronous oscillations be maintained robustly? Modeling studies suggest that synchrony can be more robustly realized in spite of heterogeneity when GABA A receptor mediated synaptic inhibition is strong [ , 53 , , 54 ]. Furthermore, network oscillations may be enhanced by subclasses of specialized neurons with distinct frequency preference due to their resonance or pacemaker biophysical properties Section II.
Finally, as we shall see in Section V, sparsely synchronized oscillations, characterized by irregular firing of individual cells, are more robust in the presence of heterogeneities. Such a model was proposed for gamma oscillations in the olfactory bulb [ , ] and in the hippocampus [ , ]. The mechanism is easy to understand as follows. An oscillatory cycle begins when fast excitation drives up neural firing in a positive feedback, until slower inhibition is recruited to eventually bring down population activity.
As the excitatory drive to interneurons wanes, the network recovers from inhibition and the next cycle starts anew, leading to rhythmic behavior [ , , , , ]. In the scenario in which a coherent oscillation is generated among a population of GABAergic cells, pyramidal neurons are presumably synchronized by inhibitory inputs to produce a population rhythm. Consistent with this scenario, an early modeling study by Lytton and Sejnowski found that IPSPs impinging on the somatic region were more effective than EPSPs on dendrites at rhythmically entraining the spike discharges of a pyramidal cell in the gamma frequency band, suggesting that perisoma-targeting basket cells play a role in phase locking pyramidal neurons [ ].
This idea was supported experimentally by Cobb et al. This is demonstrated again recently using optogenetic method to activate a subtype of genetically targeted neurons in the intact brain. Two groups reported that activating channorodopsin-2 a cation channel in fast spiking interneurons led to entrainment of pyramidal cells preferentially at gamma frequencies in mouse frontal [ ] and somatosensory [ ] cortex.
When photoactivation was administrated at different frequencies in fast spiking interneurons but not pyramidal cells, the LFP power at the stimulus frequency exhibited a pronounced peak in the gamma frequency band Fig. A Spiking activity of neurons in mouse somatosensory cortex in response to channorodopsin-2 activation of FS interneurons by repetitive light pulses at 40 Hz. LFP and raster plots of an FS cell blue and nearby regular spiking pyramidal cells recorded simultaneously RS, green.
Light pulses at 0 ms evoked FS spikes with a delay of 1—2 ms, followed by an increase in RS spiking at 17—24 ms. Power spectrum of LFP in control black, with a broad profile and with activation of FS cells by 40 Hz light pulses blue, with a sharp peak at 40 Hz. B Mean LFP power ratio in each frequency band in response to light activation of FS filled circles and RS open circles cells at those stimulus frequencies. A Top and middle panels kindly provided by Jessica Cardin, A bottom panel and B reproduced with permission from [ ]. Entrainment involves unidirectional inhibitory-to-excitatory coupling, whereas in the hippocampus [ ] and neocortex [ 85 ] interneurons and pyramidal cells are reciprocally connected.
An alternative to the interneuronal network model is that spike-to-spike synchrony emerges from the interplay between excitatory principal neurons and inhibitory interneurons. In models of interacting excitatory and inhibitory neural oscillators, two regimes were identified [ , 96 , , 62 ]. In the first regime, interneurons are sufficiently driven by external inputs and show a phase advance relative to excitatory neurons. The oscillation frequency is comparable to single neuron firing rates.
Although mutual excitation between principal neurons cannot by itself produce synchrony, it enhances oscillation generated by the feedback inhibition through amplifying excitatory neural population activity. A mathematical analysis found that the second regime is less sensitive to heterogeneity hence more robust than the first regime [ ].
In a local cortical circuit of the intact brain, both the interneuronal network mechanism and the excitatory-inhibitory feedback loop mechanism are present, and one or the other may be more predominant depending on the relative strengths of the two [ , , 96 , ]. To some extent these two mechanisms may be redundant to ensure the robustness of a synchronous rhythm. However, they are not likely to behave exactly the same way. For example, synchrony by the excitatory-inhibitory feedback loop mechanism is actually less prominent in the presence, than in the absence, of the inhibitory-to-inhibitory coupling between GABAergic neurons [ , , ].
This is because I-to-I coupling reduces the effective time constant of the inhibitory neural population, an accelerated dynamics of interneurons runs counter to a significant delay of feedback inhibition that is required in the excitatory-inhibitory loop scenario for rhythmogenesis [ ]. More importantly, in cortical circuits where neuronal activity is highly irregular, the interneuronal network and the excitatory-inhibitory loop give rise to synchronous oscillations in different frequency bands, which potentially are predominant in different brain states or serving different functions see Section V.
Short-term synaptic plasticity induces temporal filtering of presynaptic inputs, with the characteristic frequency dependence determined by the relative strength and time constants of facilitation and depression [ , , ]. At a synaptic connection exhibiting both facilitation and depression, the release probability of transmitters hence the synaptic response per presynaptic spike could be a non-monotonic function of the presynaptic firing frequency: Synaptic filtering may contribute to frequency- dependent neural responses, often in interplay with presynaptic burst firing [ , , , ].
Its role in synchronous oscillations awaits to be fully elucidated in the future. As indicated above, combining fast positive feedback and slow negative feedback offers a general recipes for rhythmogenesis. Delayed negative feedback [ ] is a versatile mechanism that can be realized, in diverse forms and on multiple timescales, in neural circuits without synaptic inhibition.
A common negative feedback process at the single cell level is spike-rate adaptation of pyramidal neurons, originating from depolarization induced activations of voltage-gated and calcium-gated potassium currents I M and I KCa , which in turn reduce the neuronal excitability. Another slow negative feedback for generating rhythmic patterns is short-term synaptic depression of recurrent synapses onto excitatory neurons [ , , , , , ], or short-term facilitation of excitatory synapses onto inhibitory interneurons [ ]. In these cases, the biophysical time constants of facilitation and depression, ranging from several hundreds of milliseconds to a few seconds, set the scale of network oscillation periodicity.
In the Down state neurons are hyperpolarized and largely silent, the activity-dependent negative feedback processes decay away, and the network recovers leading to the onset of the next Up state [ ]. The dynamics of Up and Down states vary greatly: A network model [ ] of interconnected pyramidal and interneuron populations, endowed with this cellular mechanism, reproduced salient in vivo and in vitro observations, including a gradual increase in the input resistance of pyramidal cells over the course of an Up state [ ] and a maintained balance between synaptic excitation and inhibition in spite of changes in the firing rate [ , , , ].
A more recent study showed that the same model exhibits beta and gamma frequency oscillations in the Up state similar to those observed in vivo in awake animals. This result is in line with the increasing evidence that Up states of slow oscillations during non-REM sleep resemble closely to the activated state of wakefulness in characteristics ranging from EEG or LFP patterns and spatial correlation length, to membrane fluctuations of single cells [ , , , , ] reviewed in [ , ]. Electrical synapses are gap junction mediated connections between neurons. In the mammalian brain, neuronal gap junctions are prevalent during postnatal development, and remain present in adulthood at least in certain classes of neurons, such as parvalbumin-expressing GABAergic cells [ , , ].
A direct demonstration of gap junctions requires intracellular recording simultaneously from two neurons, by depolarizing or hyperpolarizing one neuron with current injection and measuring the resulting change in the membrane potential of another neuron. Such experiments became possible only recently with the advance of new techniques, like infrared-differential interference contrast optics that allows the experimenter to visually identify recorded neurons.
Remarkably, it was found that gap junctions almost exclusively connect GABAergic interneurons within the same class. For example, perisomatic-targeting, parvalbumin-expressing FS cells are connected among themselves by gap junctions; so are dendrite-targeting, somatostatin-expressing LTS cells, but electrical synapses are rare between mixed-type FS-LTS cell pairs [ , ].
So far, at least five such distinct networks of electrically coupled GABAergic interneurons have been identified for a review see [ ].
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Compared to chemical synapses, electrical coupling has properties that seem especially desirable for subserving neural synchronization: This mechanism of intra-population coordination may be especially important for some cell types, such as LTS interneurons which exhibit a paucity of inhibitory synapses among themselves [ ]. Electrical synapses have been suggested to promote oscillatory activity [ , ] and contribute to coincidence detection in inhibitory neurons [ ]. Computational modeling studies generally found that sufficiently strong electrical synapses encourage synchrony [ , , , , , , ], but can also have desynchronizing effects depending on the interplay with inhibitory synapses and the intrinsic membrane properties of the constituent cells [ , , ].
A pair of FS interneurons or LTS neurons coupled by gap junctions showed similar phase-locking behavior, despite their differences in the biophysical properties. Interestingly, FS and LTS neurons are synchronized preferentially in the gamma and beta frequency bands, respectively [ ]. Therefore, electrical coupling is spatially very localized, like diffusion in physical and chemical systems. To experimentally assess the role of electrical synapses in brain rhythms, one must be able to induce a disruption of gap junction communications and examine its impact on a network oscillation.
Presently there are no drugs that block gap junctions with a high degree of selectivity, but genetic manipulations have proven a powerful tool. Connexin 36 is the predominant gap junction protein expressed in neurons of the mammalian brain [ ]. Both in vitro [ ] and in vivo [ ] studies showed that in mice, connexin 36 knockout only produced a reduction of the field potential gamma power or the degree of network synchronization , without affecting the oscillation frequency.
Given that electrical synapses are much faster than chemical synapses, they may be especially important for synchronizing the ultrafast — Hz sharp wave ripples. An in vitro experiment supports a critical involvement of gap junctions in the generation of sharp wave ripples [ ].