Π”ΠΈΠΏΠ»ΠΎΠΌ, курсовая, ΠΊΠΎΠ½Ρ‚Ρ€ΠΎΠ»ΡŒΠ½Π°Ρ Ρ€Π°Π±ΠΎΡ‚Π°
ΠŸΠΎΠΌΠΎΡ‰ΡŒ Π² написании студСнчСских Ρ€Π°Π±ΠΎΡ‚

ЭлСктрофизиологичСскиС коррСляты Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΡ обучСния ΠΈ памяти Ρƒ старых ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ…

Π”ΠΈΡΡΠ΅Ρ€Ρ‚Π°Ρ†ΠΈΡΠŸΠΎΠΌΠΎΡ‰ΡŒ Π² Π½Π°ΠΏΠΈΡΠ°Π½ΠΈΠΈΠ£Π·Π½Π°Ρ‚ΡŒ ΡΡ‚ΠΎΠΈΠΌΠΎΡΡ‚ΡŒΠΌΠΎΠ΅ΠΉ Ρ€Π°Π±ΠΎΡ‚Ρ‹

На Π΄Π°Π½Π½Ρ‹ΠΉ ΠΌΠΎΠΌΠ΅Π½Ρ‚ Π±ΠΎΠ»ΡŒΡˆΠΈΠ½ΡΡ‚Π²ΠΎ исслСдований Π² ΠΎΠ±Π»Π°ΡΡ‚ΠΈ памяти ΠΈ ΡΡ‚арСния сфокусировано Π½Π° Ρ€Π°Π±ΠΎΡ‚Π΅ Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ°, Π΅Π³ΠΎ Π°Ρ„Ρ„Π΅Ρ€Π΅Π½Ρ‚Π½Ρ‹Ρ… ΠΈ ΡΡ„Ρ„Π΅Ρ€Π΅Π½Ρ‚Π½Ρ‹Ρ… связях. Наряду с ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡΠΌΠΈ Π² ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠΈ, молСкулярной Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ Π±ΠΈΠΎΡ…ΠΈΠΌΠΈΠΈ сущСствуСт ΡˆΠΈΡ€ΠΎΠΊΠΈΠΉ спСктр изучСния Π½Π΅ΠΉΡ€ΠΎΠ½Π°Π»ΡŒΠ½ΠΎΠΉ пластичности ΠΌΠΎΠ·Π³Π° Π² ΠΏΡ€ΠΎΡ†Π΅ΡΡΠ΅ старСния. Богласно соврСмСнным прСдставлСниям, Π΄Π²Π° основных ΠΊΠ»Π΅Ρ‚ΠΎΡ‡Π½Ρ‹Ρ… ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠ° Π²ΠΎΠ²Π»Π΅Ρ‡Π΅Π½Ρ‹ Π² ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΠ΅… Π§ΠΈΡ‚Π°Ρ‚ΡŒ Π΅Ρ‰Ρ‘ >

Π‘ΠΎΠ΄Π΅Ρ€ΠΆΠ°Π½ΠΈΠ΅

  • 1. Π’Π’Π•Π”Π•ΠΠ˜Π•
  • 2. ΠžΠ‘Π—ΠžΠ  Π›Π˜Π’Π•Π ΠΠ’Π£Π Π«
    • 2. 1. ΠœΠΎΡ€Ρ„ΠΎΠ»ΠΎΠ³ΠΈΡ‡Π΅ΡΠΊΠΎΠ΅ строСниС Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ°
      • 2. 1. 1. ΠžΡΠ½ΠΎΠ²Π½Ρ‹Π΅ систСмы Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ°Π»ΡŒΠ½Ρ‹Ρ… связСй
      • 2. 1. 2. Буммарная элСктрофизиологичСская Π°ΠΊΡ‚ΠΈΠ²Π½ΠΎΡΡ‚ΡŒ Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ°
      • 2. 1. 3. ЭлСктрофизология ΠΏΠ΅Ρ€Π΅ΠΆΠΈΠ²Π°ΡŽΡ‰ΠΈΡ… срСзов Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ°
    • 2. 2. ПовСдСниС ΠΈ Π½Π΅ΠΉΡ€ΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Π΅ процСссы
      • 2. 2. 1. ВозрастныС измСнСния Π² ΠΏΡ€ΠΎΡΡ‚ранствСнной памяти Π½Π° ΠΏΡ€ΠΈΠΌΠ΅Ρ€Π΅ обучСния Π² Π²ΠΎΠ΄Π½ΠΎΠΌ Π»Π°Π±ΠΈΡ€ΠΈΠ½Ρ‚Π΅ ΠœΠΎΡ€Ρ€ΠΈΡΠ°
    • 2. 3. Π€Π΅Π½ΠΎΠΌΠ΅Π½ Π΄Π»ΠΈΡ‚Π΅Π»ΡŒΠ½ΠΎΠ³ΠΎ увСличСния синаптичСской пластичности (Π”Π‘ΠŸ)
      • 2. 3. 1. ΠœΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΡ‹ возникновСния ΠΈ ΠΏΡ€ΠΎΡΠ²Π»Π΅Π½ΠΈΡ Π”Π‘ΠŸ
      • 2. 3. 2. ΠŸΠΎΠ΄Π΄Π΅Ρ€ΠΆΠ°Π½ΠΈΠ΅ Π”Π‘ΠŸ
      • 2. 3. 3. ИзмСнСния Π² Π”Π‘П ΠΏΡ€ΠΈ старСнии
      • 2. 3. 4. Π˜Π½Π΄ΡƒΠΊΡ†ΠΈΡ Π”Π‘ΠŸ ΠΏΡ€ΠΈ старСнии
      • 2. 3. 5. ΠΠ°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠ΅ ΠΈΠ½Π΄ΡƒΠΊΡ†ΠΈΠΈ Π”Π‘ΠŸ совпадаСт с Π·Π°ΠΌΠ΅Π΄Π»Π΅Π½Π½Ρ‹ΠΌ ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΠ΅ΠΌ Ρƒ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈ both Ρ‹Ρ…
      • 2. 3. 6. ΠŸΠΎΠ΄Π΄Π΅Ρ€ΠΆΠ°Π½ΠΈΠ΅ Π”Π‘ΠŸ ΠΏΡ€ΠΈ стрСнии
    • 2. 4. VDCC-зависимая Ρ„ΠΎΡ€ΠΌΠ° Π”Π‘ΠŸ
      • 2. 4. 1. Π˜Π½Π΄ΡƒΠΊΡ†ΠΈΡ VDCC-Π”Π‘ΠŸ
      • 2. 4. 2. ИзмСнСния Π² VDCC Π”Π‘ΠŸ ΠΏΡ€ΠΈ старСнии
    • 2. 5. Π”Π»ΠΈΡ‚Π΅Π»ΡŒΠ½Π°Ρ синаптичСская дСпрСссия
    • 2. 6. БинаптичСскиС ΠΌΠΎΠ΄ΠΈΡ„ΠΈΠΊΠ°Ρ†ΠΈΠΈ ΠΏΡ€ΠΈ старСнии
    • 2. 7. МСдлСнная слСдовая гипСрполяризация

ЭлСктрофизиологичСскиС коррСляты Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΡ обучСния ΠΈ памяти Ρƒ старых ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… (Ρ€Π΅Ρ„Π΅Ρ€Π°Ρ‚, курсовая, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½Ρ‚Ρ€ΠΎΠ»ΡŒΠ½Π°Ρ)

Π’ Π½Π°ΡΡ‚оящСС врСмя ΠΎΠΊΠΎΠ»ΠΎ 15% насСлСния ΠΌΠΈΡ€Π° ΡΠΎΡΡ‚Π°Π²Π»ΡΡŽΡ‚ Π»ΠΈΡ†Π° ΠΏΠΎΠΆΠΈΠ»ΠΎΠ³ΠΎ ΠΈ ΡΡ‚арчСского возраста. По ΠΏΡ€ΠΎΠ³Π½ΠΎΠ·Π°ΠΌ, ΠΊ 2020 Π³ΠΎΠ΄Ρƒ ΠΈΡ… Ρ‡ΠΈΡΠ»Π΅Π½Π½ΠΎΡΡ‚ΡŒ возрастСт Π²Π΄Π²ΠΎΠ΅. Π’ Ρ€Π°Π·Π²ΠΈΡ‚Ρ‹Ρ… странах Π»ΠΈΡ†Π° ΠΏΠΎΠΆΠΈΠ»ΠΎΠ³ΠΎ возраста ΠΌΠΎΠ³ΡƒΡ‚ ΡΠΎΡΡ‚Π°Π²ΠΈΡ‚ΡŒ ΠΎΡΠ½ΠΎΠ²Π½ΡƒΡŽ Ρ‡Π°ΡΡ‚ΡŒ насСлСния. Π‘Ρ‚Π°Ρ€Π΅Π½ΠΈΠ΅ ΠΎΡ€Π³Π°Π½ΠΈΠ·ΠΌΠ° сопровоТдаСтся Ρ†Π΅Π»Ρ‹ΠΌ рядом патологичСских растройств, Ρ‚Π°ΠΊΠΈΡ… ΠΊΠ°ΠΊ болСзнь ΠΠ»ΡŒΡ†Π³Π΅ΠΉΠΌΠ΅Ρ€Π°, болСзнь ΠŸΠ°Ρ€ΠΊΠΈΠ½ΡΠΎΠ½Π° ΠΈ Π΄Π΅ΠΌΠ΅Π½Ρ†ΠΈΡ. Π’ ΠΏΠΎΠ΄Π°Π²Π»ΡΡŽΡ‰Π΅ΠΌ Π±ΠΎΠ»ΡŒΡˆΠΈΠ½ΡΡ‚Π²Π΅ случаСв, Π΄Π°ΠΆΠ΅ ΠΏΡ€ΠΈ отсутствии Π²Ρ‹Ρ€Π°ΠΆΠ΅Π½Π½ΠΎΠΉ ΠΏΠ°Ρ‚ΠΎΠ»ΠΎΠ³ΠΈΠΈ, отмСчаСтся сниТСниС Π²Ρ‹ΡΡˆΠΈΡ… ΠΊΠΎΠ³Π½ΠΈΡ‚ΠΈΠ²Π½Ρ‹Ρ… Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΉ, извСстноС ΠΊΠ°ΠΊ ΡƒΠΌΠ΅Ρ€Π΅Π½Π½Ρ‹Π΅ ΠΊΠΎΠ³Π½ΠΈΡ‚ΠΈΠ²Π½Ρ‹Π΅ растройства (mild cognitive impairments, MCI). ΠŸΡ€Π΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡ‚Π΅Π»ΡŒΠ½ΠΎ, ΡƒΠΌΠ΅Ρ€Π΅Π½Π½Ρ‹Π΅ ΠΊΠΎΠ³Π½ΠΈΡ‚ΠΈΠ²Π½Ρ‹Π΅ растройства ΡΠ²Π»ΡΡŽΡ‚ΡΡ ΠΏΡ€Π΅Π΄ΡˆΠ΅ΡΡ‚Π²Π΅Π½Π½ΠΈΠΊΠ°ΠΌΠΈ Π±ΠΎΠ»Π΅Π΅ ΡΠ΅Ρ€ΡŒΠ΅Π·Π½ΠΎΠΉ ΠΏΠ°Ρ‚Π°Π»ΠΎΠ³ΠΈΠΈ. Π’ ΡΡ‚ΠΎΠΉ связи, ΠΈΠ·ΡƒΡ‡Π΅Π½ΠΈΠ΅ ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠΎΠ² памяти, Π° Ρ‚Π°ΠΊΠΆΠ΅ поиск ΠΈ ΡΠΎΠ·Π΄Π°Π½ΠΈΠ΅ фармакологичСских ΠΏΡ€Π΅ΠΏΠ°Ρ€Π°Ρ‚ΠΎΠ², ΠΎΠ±Π»Π°Π΄Π°ΡŽΡ‰ΠΈΡ… ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡ‚ΡŒΡŽ Π·Π°ΠΌΠ΅Π΄Π»ΡΡ‚ΡŒ/ΠΏΡ€Π΅Π΄ΠΎΡ‚Π²Ρ€Π°Ρ‰Π°Ρ‚ΡŒ Ρ€Π°Π·Π²ΠΈΡ‚ΠΈΠ΅ Π±ΠΎΠ»Π΅Π·Π½ΠΈ ΠΈ ΡƒΠ»ΡƒΡ‡ΡˆΠ°Ρ‚ΡŒ ΠΏΠ°ΠΌΡΡ‚ΡŒ Π² ΡΠ»ΡƒΡ‡Π°Π΅ ΡƒΠΌΠ΅Ρ€Π΅Π½Π½Ρ‹Ρ… ΠΊΠΎΠ³Π½ΠΈΡ‚ΠΈΠ²Π½Ρ‹Ρ… расстройств, являСтся ΠΎΠ΄Π½ΠΎΠΉ ΠΈΠ· Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ Π°ΠΊΡ‚ΡƒΠ°Π»ΡŒΠ½Ρ‹Ρ… ΠΏΡ€ΠΎΠ±Π»Π΅ΠΌ Π½Π΅ΠΉΡ€ΠΎΠ±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ Π½Π΅ΠΉΡ€ΠΎΡ„Π°Ρ€ΠΌΠ°ΠΊΠΎΠ»ΠΎΠ³ΠΈΠΈ.

Π’ ΠΏΡ€ΠΎΡ†Π΅ΡΡΠ΅ старСния измСняСтся Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½ΠΈΡ€ΠΎΠ²Π°Π½ΠΈΠ΅ ΠΌΠΎΠ·Π³Π°, обусловлСноС ΠΊΠ°ΠΊ Ρ€Π°Π·Π²ΠΈΡ‚ΠΈΠ΅ΠΌ Π΄Π΅Π³Π΅Π½Π΅Ρ€Π°Ρ‚ΠΈΠ²Π½Ρ‹Ρ… процСссов, Ρ‚Π°ΠΊ ΠΈ ΠΈΠ½Ρ‚СнсификациСй компСнсаторных явлСний. ΠŸΡ€ΠΈ этом происходит ряд морфологичСских ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠΉ Π² Ρ†Π΅Π½Ρ‚Ρ€Π°Π»ΡŒΠ½ΠΎΠΉ Π½Π΅Ρ€Π²Π½ΠΎΠΉ систСмС, ΠΊΠΎΡ‚ΠΎΡ€Ρ‹Π΅ Π² ΠΏΠ΅Ρ€Π²ΡƒΡŽ ΠΎΡ‡Π΅Ρ€Π΅Π΄ΡŒ приводят ΠΊ ΠΎΡΠ»Π°Π±Π»Π΅Π½ΠΈΡŽ памяти ΠΈ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡ‚ΠΈ ΠΎΠ±ΡƒΡ‡Π°Ρ‚ΡŒΡΡ. Одна ΠΈΠ· ΠΎΡΠ½ΠΎΠ²Π½Ρ‹Ρ… ΠΏΡ€ΠΎΠ±Π»Π΅ΠΌ, стоящих ΠΏΠ΅Ρ€Π΅Π΄ ΡƒΡ‡Π΅Π½Ρ‹ΠΌΠΈ, это Π½Π°ΠΉΡ‚ΠΈ Ρ‡Π΅Ρ‚ΠΊΠΈΠ΅ ΠΊΡ€ΠΈΡ‚Π΅Ρ€ΠΈΠΈ, Ρ€Π°Π·Π΄Π΅Π»ΡΡŽΡ‰ΠΈΠ΅ Π½ΠΎΡ€ΠΌΠ°Π»ΡŒΠ½ΠΎΠ΅ старСниС ΠΎΡ‚ ΠΏΠ°Ρ‚ΠΎΠ»ΠΎΠ³ΠΈΠΈ. Для Ρ€Π΅ΡˆΠ΅Π½ΠΈΡ этой Π·Π°Π΄Π°Ρ‡ΠΈ ΡƒΡ‡Π΅Π½Ρ‹Π΅ ΡˆΠΈΡ€ΠΎΠΊΠΎ ΠΈΡΠΏΠΎΠ»ΡŒΠ·ΡƒΡŽΡ‚ Π»Π°Π±ΠΎΡ€Π°Ρ‚ΠΎΡ€Π½Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… Π² ΠΊΠ°Ρ‡Π΅ΡΡ‚Π²Π΅ ΠΌΠΎΠ΄Π΅Π»Π΅ΠΉ ΠΊΠ°ΠΊ для нСйрологичСских Π±ΠΎΠ»Π΅Π·Π½Π΅ΠΉ, Ρ‚Π°ΠΊ ΠΈ Π΄Π»Ρ Π½ΠΎΡ€ΠΌΠ°Π»ΡŒΠ½ΠΎΠ³ΠΎ старСния. Π”Π°Π½Π½Ρ‹Π΅, ΠΏΠΎΠ»ΡƒΡ‡Π΅Π½Ρ‹Π΅ Π² Ρ€Π΅Π·ΡƒΠ»ΡŒΡ‚Π°Ρ‚Π΅ экспСримСнтов Π½Π° Π»Π°Π±ΠΎΡ€Π°Ρ‚ΠΎΡ€Π½Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… ΠΎΠΊΠ°Π·Ρ‹Π²Π°ΡŽΡ‚ΡΡ ΠΏΡ€ΠΈΠΌΠ΅Π½ΠΈΠΌΡ‹ для характСристики физиологичСских, гСнСтичСских ΠΈ ΠΌΠΎΠ»Π΅ΠΊΡƒΠ»ΡΡ€Π½Ρ‹Ρ… ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠΉ, происходящих Ρƒ Π»ΡŽΠ΄Π΅ΠΉ Π²ΠΎ Π²Ρ€Π΅ΠΌΡ старСния. Π’Π°ΠΊ ΠΊΠ°ΠΊ Π² ΠΎΡ‚Π»ΠΈΡ‡ΠΈΠ΅ ΠΎΡ‚ Π»ΡŽΠ΄Π΅ΠΉ, Π»Π°Π±ΠΎΡ€Π°Ρ‚ΠΎΡ€Π½Ρ‹Π΅ ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Π΅ Π½Π΅ Ρ€Π°Π·Π²ΠΈΠ²Π°ΡŽΡ‚ Π½Π΅ΠΉΡ€ΠΎΠ΄Π΅Π³Π΅Π½Π΅Ρ€Π°Ρ‚ΠΈΠ²Π½Ρ‹Π΅ Π±ΠΎΠ»Π΅Π·Π½ΠΈ, Ρ‚ΠΈΠΏΠ° Π±ΠΎΠ»Π΅Π·Π½ΠΈ ΠΠ»ΡŒΡ†Π³Π΅ΠΉΠΌΠ΅Ρ€Π°, Ρ‚ΠΎ Π΅ΡΡ‚ΡŒ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡ‚ΡŒ изучСния Π½ΠΎΡ€ΠΌΠ°Π»ΡŒΠ½ΠΎΠ³ΠΎ старСния Π² Ρ‡ΠΈΡΡ‚ΠΎΠΌ Π²ΠΈΠ΄Π΅. Однако ΠΎΠ΄Π½ΠΎΠΉ ΠΈΠ· ΠΏΡ€ΠΎΠ±Π»Π΅ΠΌ, Π²ΠΎΠ·Π½ΠΈΠΊΠ°ΡŽΡ‰ΠΈΡ… ΠΏΡ€ΠΈ Ρ€Π°Π±ΠΎΡ‚Π΅ со ΡΡ‚Π°Ρ€Ρ‹ΠΌΠΈ ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹ΠΌΠΈ являСтся Ρ‚ΠΎ, Ρ‡Ρ‚ΠΎ Ρ„Π΅Π½ΠΎΡ‚ΠΈΠΏ Π½ΠΎΡ€ΠΌΠ°Π»ΡŒΠ½ΠΎΠ³ΠΎ старСния ΠΈΠΌΠ΅Π΅Ρ‚ Π±ΠΎΠ»ΡŒΡˆΡƒΡŽ Π²Π°Ρ€ΠΈΠ°Π±Π΅Π»ΡŒΠ½ΠΎΡΡ‚ΡŒ. Π­Ρ‚Π° Π²Π°Ρ€ΠΈΠ°Π±Π΅Π»ΡŒΠ½ΠΎΡΡ‚ΡŒ часто Π½Π°Π±Π»ΡŽΠ΄Π°Π΅Ρ‚ΡΡ ΠΊΠ°ΠΊ ΠΌΠ΅ΠΆΠ΄Ρƒ линиями ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ…, Ρ‚Π°ΠΊ ΠΈ Π²Π½ΡƒΡ‚Ρ€ΠΈ ΠΎΠ΄Π½ΠΎΠΉ Π»ΠΈΠ½ΠΈΠΈ, ΠΏΠΎΡΠΊΠΎΠ»ΡŒΠΊΡƒ процСсс старСния опосрСдован Ρ†Π΅Π»Ρ‹ΠΌ рядом гСнСтичСских Ρ„Π°ΠΊΡ‚ΠΎΡ€ΠΎΠ², ΠΏΡ€ΠΎΡΠ²Π»ΡΡŽΡ‰ΠΈΡ…ΡΡ Π² Π²ΠΈΠ΄Π΅ Ρ€Π°Π·Π»ΠΈΡ‡Π½Ρ‹Ρ… Ρ„Π΅Π½ΠΎΡ‚ΠΈΠΏΠΎΠ². По ΡΡ‚ΠΎΠΉ ΠΏΡ€ΠΈΡ‡ΠΈΠ½Π΅ ΠΏΡ€ΠΈ исслСдовании ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠΎΠ² старСния Π² ΠΏΠΎΡΠ»Π΅Π΄Π½Π΅Π΅ врСмя стали ΠΈΡΠΏΠΎΠ»ΡŒΠ·ΠΎΠ²Π°Ρ‚ΡŒ ΠΌΡ‹ΡˆΠ΅ΠΉ с Π³Π΅Π½Π΅Ρ‚ичСскими модификациями. ИспользованиС ΠΌΡ‹ΡˆΠ΅ΠΉ с Π½Π°ΠΏΡ€Π°Π²Π»Π΅Π½Π½ΠΎΠΉ ΠΌΡƒΡ‚Π°Ρ†ΠΈΠ΅ΠΉ Π³Π΅Π½ΠΎΠ², ΠΊΠ°ΠΊ модСль для ΠΎΠΏΡ€Π΅Π΄Π΅Π»Π΅Π½Π½Ρ‹Ρ… гСнСтичСских манифСстаций Π² Π½Π°ΡΡ‚оящСС врСмя ΡˆΠΈΡ€ΠΎΠΊΠΎ примСняСтся Π² ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡΡ… Π±ΠΎΠ»Π΅Π·Π½ΠΈ ΠΠ»ΡŒΡ†Π³Π΅ΠΉΠΌΠ΅Ρ€Π°, ΡΠΈΠ½ΠΈΠ»ΡŒΠ½Ρ‹Ρ… расстройств ΠΈ Π±ΠΎΠ»Π΅Π·Π½ΠΈ ΠŸΠ°Ρ€ΠΊΠΈΠ½ΡΠΎΠ½Π°.

На Π΄Π°Π½Π½Ρ‹ΠΉ ΠΌΠΎΠΌΠ΅Π½Ρ‚ Π±ΠΎΠ»ΡŒΡˆΠΈΠ½ΡΡ‚Π²ΠΎ исслСдований Π² ΠΎΠ±Π»Π°ΡΡ‚ΠΈ памяти ΠΈ ΡΡ‚арСния сфокусировано Π½Π° Ρ€Π°Π±ΠΎΡ‚Π΅ Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ°, Π΅Π³ΠΎ Π°Ρ„Ρ„Π΅Ρ€Π΅Π½Ρ‚Π½Ρ‹Ρ… ΠΈ ΡΡ„Ρ„Π΅Ρ€Π΅Π½Ρ‚Π½Ρ‹Ρ… связях. Наряду с ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡΠΌΠΈ Π² ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠΈ, молСкулярной Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ Π±ΠΈΠΎΡ…ΠΈΠΌΠΈΠΈ сущСствуСт ΡˆΠΈΡ€ΠΎΠΊΠΈΠΉ спСктр изучСния Π½Π΅ΠΉΡ€ΠΎΠ½Π°Π»ΡŒΠ½ΠΎΠΉ пластичности ΠΌΠΎΠ·Π³Π° Π² ΠΏΡ€ΠΎΡ†Π΅ΡΡΠ΅ старСния. Богласно соврСмСнным прСдставлСниям, Π΄Π²Π° основных ΠΊΠ»Π΅Ρ‚ΠΎΡ‡Π½Ρ‹Ρ… ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠ° Π²ΠΎΠ²Π»Π΅Ρ‡Π΅Π½Ρ‹ Π² ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΠ΅ ΠΈ ΠΏΠ°ΠΌΡΡ‚ΡŒ. ΠŸΠ΅Ρ€Π²Ρ‹ΠΉ, Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ Π΄Π΅Ρ‚Π°Π»ΡŒΠ½ΠΎ ΠΈΠ·ΡƒΡ‡Π΅Π½Π½Ρ‹ΠΉ, это Π΄Π»ΠΈΡ‚Π΅Π»ΡŒΠ½Π°Ρ синаптичСская потСнциация (Π”Π‘ΠŸ), ΠΏΡ€Π΅Π΄ΡΡ‚Π°Π²Π»ΡΡŽΡ‰Π°Ρ собой ΠΏΡ€ΠΎΠ΄ΠΎΠ»ΠΆΠΈΡ‚Π΅Π»ΡŒΠ½ΠΎΠ΅ ΡƒΠ²Π΅Π»ΠΈΡ‡Π΅Π½ΠΈΠ΅ эффСктивности синаптичСской ΠΏΠ΅Ρ€Π΅Π΄Π°Ρ‡ΠΈ Π² ΠΎΡ‚Π²Π΅Ρ‚ Π½Π° ΠΊΡ€Π°Ρ‚ΠΊΠΎΠ²Ρ€Π΅ΠΌΠ΅Π½Π½ΡƒΡŽ Π²Ρ‹ΡΠΎΠΊΠΎΡ‡Π°ΡΡ‚ΠΎΡ‚Π½ΡƒΡŽ ΡΡ‚ΠΈΠΌΡƒΠ»ΡΡ†ΠΈΡŽ (Bliss, 2003). Π’Ρ‚ΠΎΡ€ΠΎΠΉ, ΠΌΠ΅Π½Π΅Π΅ ΠΈΠ·ΡƒΡ‡Π΅Π½ΠΈΠΉ, это Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Π΅ измСнСния возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ², Ρ€Π΅Π³ΡƒΠ»ΠΈΡ€ΡƒΠ΅ΠΌΡ‹Π΅ ΠΌΠ΅Π΄Π»Π΅Π½Π½ΠΎΠΉ слСдовой гипСрполяризациСй (ΠœΠ‘Π“) Π½Π° ΡƒΡ€ΠΎΠ²Π½Π΅ Ρ‚Π΅Π»Π° ΠΊΠ»Π΅Ρ‚ΠΊΠΈ (Daudal and Debanne, 2003; Disterhoft et al., 2004; Giese et al., 2001; Murphy et al., 2004; Sourdet et al., 2003). Оба ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠ° Π²ΠΎΠ²Π»Π΅Ρ‡Π΅Π½Ρ‹ Π² ΠΎΠ±Ρ€Π°Π·ΠΎΠ²Π°Π½ΠΈΠ΅/усилСниС Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Ρ… связСй ΠΌΠ΅ΠΆΠ΄Ρƒ Π½Π΅ΠΉΡ€ΠΎΠ½Π°ΠΌΠΈ Π»ΠΈΠ±ΠΎ Π·Π° ΡΡ‡Π΅Ρ‚ Π΄Π»ΠΈΡ‚Π΅Π»ΡŒΠ½ΠΎΠ³ΠΎ увСличСния силы синаптичСских связСй, Π»ΠΈΠ±ΠΎ Π·Π° ΡΡ‡Π΅Ρ‚ ΠΏΡ€ΠΎΠ΄ΠΎΠ»ΠΆΠΈΡ‚Π΅Π»ΡŒΠ½ΠΎΠ³ΠΎ возрастания возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ². Π˜Π·Π²Π΅ΡΡ‚Π½ΠΎ, Ρ‡Ρ‚ΠΎ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠ΅ любого ΠΈΠ· ΡΡ‚ΠΈΡ… процСссов ΠΎΡ‚Ρ€ΠΈΡ†Π°Ρ‚Π΅Π»ΡŒΠ½ΠΎ влияСт Π½Π° ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΠ΅ ΠΈ ΠΏΠ°ΠΌΡΡ‚ΡŒ Π»Π°Π±ΠΎΡ€Π°Ρ‚ΠΎΡ€Π½Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… (Disterhoft et al., 2004; Giese et al., 1998). Π’Π°ΠΊΠΈΠΌ ΠΎΠ±Ρ€Π°Π·ΠΎΠΌ, Ρ€Π΅Π·ΡƒΠ»ΡŒΡ‚Π°Ρ‚Ρ‹ Π΄Π°Π½Π½ΠΎΠΉ Ρ€Π°Π±ΠΎΡ‚Ρ‹ ΠΌΠΎΠ³ΡƒΡ‚ Π±Ρ‹Ρ‚ΡŒ ΠΏΠΎΠ»Π΅Π·Π½Ρ‹ для Π±ΠΎΠ»Π΅Π΅ Π³Π»ΡƒΠ±ΠΎΠΊΠΎΠ³ΠΎ понимания ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠΎΠ² Π»Π΅ΠΆΠ°Ρ‰ΠΈΡ… Π² ΠΎΡΠ½ΠΎΠ²Π΅ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠΉ обучСния ΠΈ ΠΏΠ°ΠΌΡΡ‚ΠΈ ΠΏΡ€ΠΈ старСнии.

Π¦Π΅Π»ΠΈ ΠΈ Π·Π°Π΄Π°Ρ‡ΠΈ исслСдования: ЦСлью Π΄Π°Π½Π½ΠΎΠ³ΠΎ исслСдования Π±Ρ‹Π»ΠΎ ΠΈΠ·ΡƒΡ‡Π΅Π½ΠΈΠ΅ элСкрофизиологичСских ΠΏΠΎΠΊΠ°Π·Π°Ρ‚Π΅Π»Π΅ΠΉ старСния Π² ΠΌΠ΅Ρ‚ΠΎΠ΄ΠΈΠΊΠ΅ in vitro, Π° Ρ‚Π°ΠΊΠΆΠ΅ ΠΈΠ·ΡƒΡ‡Π΅Π½ΠΈΠ΅ взаимосвязи ΠΌΠ΅ΠΆΠ΄Ρƒ ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΠ΅ΠΌ ΠΈ ΡΠ»Π΅ΠΊΡ‚рофизиологичСскими показатСлями. ΠšΠΎΠ½ΠΊΡ€Π΅Ρ‚Π½Ρ‹Π΅ Π·Π°Π΄Π°Ρ‡ΠΈ Π΄Π°Π½Π½ΠΎΠ³ΠΎ исслСдования состояли Π² ΡΠ»Π΅Π΄ΡƒΡŽΡ‰Π΅ΠΌ:

1. ΠΠ΄Π°ΠΏΡ‚ΠΈΡ€ΠΎΠ²Π°Ρ‚ΡŒ ΠΌΠ΅Ρ‚ΠΎΠ΄ΠΈΠΊΡƒ ΠΏΠ΅Ρ€Π΅ΠΆΠΈΠ²Π°ΡŽΡ‰ΠΈΡ… срСзов Π³ΠΈΠΏΠΏΠΎΠΊΠ°ΠΌΠΏΠ° для Ρ€Π°Π±ΠΎΡ‚Ρ‹ со ΡΡ‚Π°Ρ€Ρ‹ΠΌΠΈ ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹ΠΌΠΈ.

2. ΠžΠΏΡ€Π΅Π΄Π΅Π»ΠΈΡ‚ΡŒ Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Π΅ измСнСния ΠœΠ‘Π“ ΠΏΠΎΠ΄ дСйствиСм Π²Π½ΡƒΡ‚Ρ€ΠΈΠΊΠ»Π΅Ρ‚ΠΎΡ‡Π½ΠΎΠΉ стимуляции ΠΈ Ρ„армакологичСских Π°Π³Π΅Π½Ρ‚ΠΎΠ².

3. ΠžΠΏΠΈΡΠ°Ρ‚ΡŒ измСнСния ΠœΠ‘Π“ Ρƒ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… ΠΈ ΠΏΡ€ΠΎΠ²Π΅Ρ€ΠΈΡ‚ΡŒ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡ‚ΡŒ фармакологичСской компСнсации Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠΉ.

4. Π‘Ρ€Π°Π²Π½ΠΈΡ‚ΡŒ Π£Π­Π‘Π‘ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ‚ Π”Π‘ΠŸ Ρƒ ΠΌΠΎΠ»ΠΎΠ΄Ρ‹Ρ… ΠΈ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… ΠΈ ΠΏΡ€ΠΎΠ²Π΅Ρ€ΠΈΡ‚ΡŒ дСйствиС Π±Π»ΠΎΠΊΠ°Ρ‚ΠΎΡ€ΠΎΠ² ΠΊΠ°Π»ΡŒΡ†ΠΈΠ΅Π²Ρ‹Ρ… ΠΊΠ°Π½Π°Π»ΠΎΠ² Π½Π° Π£Π‘Π‘Π‘ ΠΈ Π«ΠœΠ­ΠΠ·Π°Π²ΠΈΡΠΈΠΌΡ‹ΠΉ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ‚Ρ‹ Π”Π‘ΠŸ.

5. ΠžΠΏΡ€Π΅Π΄Π΅Π»ΠΈΡ‚ΡŒ взаимосвязь ΠΌΠ΅ΠΆΠ΄Ρƒ элСктрофизиологичСскими показатСлями ΠΈ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠ΅ΠΌ обучСния Ρƒ Π»Π°Π±ΠΎΡ€Π°Ρ‚ΠΎΡ€Π½Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ….

6. ΠžΠΏΡ€Π΅Π΄Π΅Π»ΠΈΡ‚ΡŒ Ρ€ΠΎΠ»ΡŒ Π”Π‘ΠŸ ΠΈ ΠœΠ‘Π“ Π² ΠΏΡ€ΠΎΡ†Π΅ΡΡΠ΅ обучСния.

Научная Π½ΠΎΠ²ΠΈΠ·Π½Π° ΠΈ ΠΏΡ€Π°ΠΊΡ‚ичСская Π·Π½Π°Ρ‡ΠΈΠΌΠΎΡΡ‚ΡŒ. Π’ Π½Π°ΡΡ‚оящСй Ρ€Π°Π±ΠΎΡ‚Π΅ Π²ΠΏΠ΅Ρ€Π²Ρ‹Π΅ Π±Ρ‹Π»ΠΈ описаны Π΄Π»ΠΈΡ‚Π΅Π»ΡŒΠ½Ρ‹Π΅ Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Π΅ измСнСния возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ². Π‘Ρ‹Π»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, Ρ‡Ρ‚ΠΎ этот процСсс, опосрСдованный Ρ‡Π΅Ρ€Π΅Π· ΠœΠ‘Π“ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ Ρƒ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ…. Π‘Ρ‹Π»Π° ΠΏΠΎΠΊΠ°Π·Π°Π½Π° прямая Π·Π°Π²ΠΈΡΠΈΠΌΠΎΡΡ‚ΡŒ ΠΌΠ΅ΠΆΠ΄Ρƒ ΡƒΡ€ΠΎΠ²Π½Π΅ΠΌ Ρ†ΠΠœΠ€ Π² Π½Π΅ΠΉΡ€ΠΎΠ½Π΅ ΠΈ Π°ΠΌΠΏΠ»ΠΈΡ‚ΡƒΠ΄ΠΎΠΉ ΠœΠ‘Π“. Π’ΠΏΠ΅Ρ€Π²Ρ‹Π΅ Π±Ρ‹Π»Π° ΠΏΠΎΠΊΠ°Π·Π°Π½Π° Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡ‚ΡŒ компСнсации Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠΉ Ρƒ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… ΠΏΡ€ΠΈ использовании фармакологичСских Π°Π³Π΅Π½Ρ‚ΠΎΠ², ΠΊΠΎΡ‚ΠΎΡ€Ρ‹Π΅ ΠΏΠΎΠ²Ρ‹ΡˆΠ°ΡŽΡ‚ ΡƒΡ€ΠΎΠ²Π΅Π½ΡŒ Ρ†ΠΠœΠ€ Π² Π½Π΅ΠΉΡ€ΠΎΠ½Π΅. Π­Ρ‚ΠΎ Π΄Π°Π΅Ρ‚ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡ‚Π΅Π»ΡŒΠ½Ρ‹Π΅ свСдСния ΠΎ Π±ΠΈΠΎΡ…имичСских процСссах, ΠΊΠΎΡ‚ΠΎΡ€Ρ‹Π΅ Π²ΠΎΠ²Π»Π΅Ρ‡Π΅Π½Ρ‹ Π² ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌ рСгуляции ΠœΠ‘Π“, Π° Ρ‚Π°ΠΊΠΆΠ΅ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡ‚ΡŒ развития лСкарствСнных ΠΏΡ€Π΅ΠΏΠ°Ρ€Π°Ρ‚ΠΎΠ² для восстановлСния памяти Π² ΠΏΠΎΠΆΠΈΠ»ΠΎΠΌ возрастС.

Π”Π°Π½Π½Ρ‹Π΅, ΠΏΠΎΠ»ΡƒΡ‡Π΅Π½Π½Ρ‹Π΅ ΠΏΡ€ΠΈ сравнСнии Π£Π­Π‘Π‘ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ‚Π° Ρƒ ΠΌΠΎΠ»ΠΎΠ΄Ρ‹Ρ… ΠΈ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… ΠΏΠΎΠ΄Ρ‡Π΅Ρ€ΠΊΠΈΠ²Π°ΡŽΡ‚ Ρ€ΠΎΠ»ΡŒ ΠΊΠ°Π»ΡŒΡ†ΠΈΡ ΠΈ ΠΊΠ°Π»ΡŒΡ†ΠΈΠ΅Π²Ρ‹Ρ… ΠΊΠ°Π½Π°Π»ΠΎΠ² ΠΏΡ€ΠΈ старСнии. Π‘Ρ‹Π»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, Ρ‡Ρ‚ΠΎ Π±Π»ΠΎΠΊΠ°Ρ‚ΠΎΡ€Ρ‹ ΠΊΠ°Π»ΡŒΡ†ΠΈΠ΅Π²Ρ‹Ρ… ΠΊΠ°Π½Π°Π»ΠΎΠ² ΠΏΠΎΠ΄Π°Π²Π»ΡΡŽΡ‚ УОББ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ‚ Π”Π‘ΠŸ, ΠΊΠΎΡ‚ΠΎΡ€Ρ‹ΠΉ ΡƒΠ²Π΅Π»ΠΈΡ‡Π΅Π½ Ρƒ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ…, Ρ‡Ρ‚ΠΎ Π΄Π΅Π»Π°Π΅Ρ‚ это Π½Π°ΠΏΡ€Π°Π²Π»Π΅Π½ΠΈΠ΅ пСрспСктивным для Ρ„Π°Ρ€ΠΌΠ°ΠΊΠΎΠ»ΠΎΠ³ΠΈΠΈ старСния.

Π’ΠΏΠ΅Ρ€Π²Ρ‹Π΅ Π±Ρ‹Π»ΠΎ ΠΎΠ±Π½Π°Ρ€ΡƒΠΆΠ΅Π½ΠΎ, Ρ‡Ρ‚ΠΎ сниТСниС возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ², вслСдствии возрастания Π°ΠΌΠΏΠ»ΠΈΡ‚ΡƒΠ΄Ρ‹ ΠœΠ‘Π“ являСтся ΠΏΡ€ΠΈΡ‡ΠΈΠ½ΠΎΠΉ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΡ пространствСнного обучСния Ρƒ ΠΌΡ‹ΡˆΠ΅ΠΉ. Π”Π°Π»Π΅Π΅ ΠΏΠΎΠ΄Ρ€ΠΎΠ±Π½ΠΎ Π±Ρ‹Π»Π° исслСдована Ρ€ΠΎΠ»ΡŒ Π² ΠΏΡ€ΠΎΡΡ‚ранствСнном ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΠΈΠΈ ΠΊΠ°ΠΆΠ΄ΠΎΠ³ΠΎ ΠΈΠ· ΡΠ»Π΅ΠΊΡ‚рофизиологичСских процСссов: ΠœΠ‘Π“ ΠΈ Π”Π‘П. Π˜Π·ΡƒΡ‡Π°Ρ коррСляции элСктрофизиологичСских Π΄Π°Π½Π½Ρ‹Ρ… с ΠΊΠ°ΠΆΠ΄Ρ‹ΠΌ Π΄Π½Π΅ΠΌ обучСния ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… Π² Π²ΠΎΠ΄Π½ΠΎΠΌ Π»Π°Π±ΠΈΡ€ΠΈΠ½Ρ‚Π΅ ΠœΠΎΡ€Ρ€ΠΈΡΠ°, Π±Ρ‹Π»ΠΎ Π²ΠΏΠ΅Ρ€Π²Ρ‹Π΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, Ρ‡Ρ‚ΠΎ ΠœΠ‘Π“ ΠΈ Π”Π‘П ΡΠ²Π»ΡΡŽΡ‚ΡΡ двумя нСзависимыми ΠΈ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°Ρ‚Π΅Π»ΡŒΠ½Ρ‹ΠΌΠΈ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ‚Π°ΠΌΠΈ ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠ° обучСния. Π­Ρ‚ΠΎ ΠΏΡ€Π΅Π΄Π»Π°Π³Π°Π΅Ρ‚ Π½ΠΎΠ²Ρ‹ΠΉ ΠΏΠΎΠ΄Ρ…ΠΎΠ΄ ΠΊ ΠΈΠ·ΡƒΡ‡Π΅Π½ΠΈΡŽ ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠΎΠ² обучСния ΠΈ ΠΏΠ°ΠΌΡΡ‚ΠΈ, Ρ‚Π°ΠΊ ΠΊΠ°ΠΊ ΠΏΠΎΠ΄ΠΎΠ±Π½Ρ‹Π΅ коррСляции Π΅Ρ‰Π΅ Π½Π΅ Π±Ρ‹Π»ΠΈ описаны Π² Π½Π°ΡƒΡ‡Π½ΠΎΠΉ Π»ΠΈΡ‚Π΅Ρ€Π°Ρ‚ΡƒΡ€Π΅.

2. ΠžΠ±Π·ΠΎΡ€ Π»ΠΈΡ‚Π΅Ρ€Π°Ρ‚ΡƒΡ€Ρ‹.

Π’Ρ‹Π²ΠΎΠ΄Ρ‹.

1) Π£ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ… происходит ΡƒΠ²Π΅Π»ΠΈΡ‡Π΅Π½ΠΈΠ΅ Π°ΠΌΠΏΠ»ΠΈΡ‚ΡƒΠ΄Ρ‹ ΠœΠ‘Π“ ΠΈ ΡΠ½ΠΈΠΆΠ΅Π½ΠΈΠ΅ Π½Π΅ΠΉΡ€ΠΎΠ½Π°Π»ΡŒΠ½ΠΎΠΉ возбудимости. ΠŸΡ€Π΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡ‚Π΅Π»ΡŒΠ½ΠΎ, сниТСниС возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ² ΠΏΡ€ΠΈΠ²ΠΎΠ΄ΠΈΡ‚ ΠΊ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΡŽ ΠΈΠ»ΠΈ замСдлСнию процСссов обучСния ΠΈ ΠΏΠ°ΠΌΡΡ‚ΠΈ.

2) ΠœΠ‘Π“ ΠΌΠΎΠΆΠ΅Ρ‚ ΠΎΠΏΠΎΡΡ€Π΅Π΄ΠΎΠ²Π°Ρ‚ΡŒ Π΄Π»ΠΈΡ‚Π΅Π»ΡŒΠ½Ρ‹Π΅ Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Π΅ измСнСния возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ². ΠΠΊΡ‚ΠΈΠ²Π½ΠΎΡΡ‚ΡŒ Ρ„Π΅Ρ€ΠΌΠ΅Π½Ρ‚Π° фосфодиСстСразы 4, ΠΊΠΎΠ½Ρ‚Ρ€ΠΎΠ»ΠΈΡ€ΡƒΡŽΡ‰Π΅Π³ΠΎ ΡƒΡ€ΠΎΠ²Π΅Π½ΡŒ Ρ†ΠΠœΠ€ Π² ΠΊΠ»Π΅Ρ‚ΠΊΠ΅, слуТит Π·Π°Ρ‚Π²ΠΎΡ€Π½Ρ‹ΠΌ ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠΎΠΌ этих ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠΉ.

3) ΠŸΡ€ΠΎΡ†Π΅ΡΡ Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½ΠΎΠΉ рСгуляции возбудимости Π½Π΅ΠΉΡ€ΠΎΠ½ΠΎΠ² Π½Π°Ρ€ΡƒΡˆΠ΅Π½ Ρƒ ΡΡ‚Π°Ρ€Ρ‹Ρ… ΠΆΠΈΠ²ΠΎΡ‚Π½Ρ‹Ρ…. Π­Ρ‚ΠΎ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠ΅ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎ ΠΊΠΎΠΌΠΏΠ΅Π½ΡΠΈΡ€ΠΎΠ²Π°Ρ‚ΡŒ ΠΏΡ€ΠΈ ΠΏΠΎΠΌΠΎΡ‰ΠΈ фармакологичСской Π±Π»ΠΎΠΊΠ°Π΄Ρ‹ фосфодиСстСразы 4 Ρ€ΠΎΠ»ΠΈΠΏΡ€Π°ΠΌΠΎΠΌ.

4) ΠŸΡ€ΠΈ старСнии происходит возрастаниС VDCC ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ‚Π° Π”Π‘ΠŸ, ΠΊΠΎΡ‚ΠΎΡ€Ρ‹ΠΉ Π·Π°ΠΌΠ΅Ρ‰Π°Π΅Ρ‚ NMDA Π·Π°Π²ΠΈΡΠΈΠΌΡƒΡŽ Π”Π‘ΠŸ. Π­Ρ‚ΠΈ Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΡ Π±ΠΎΠ³ΡƒΡ‚ Π±Ρ‹Ρ‚ΡŒ компСнсированы Π±Π»ΠΎΠΊΠ°Ρ‚ΠΎΡ€Π°ΠΌΠΈ VDCC ΠΊΠ°Π½Π°Π»ΠΎΠ² (Π½Π΅Ρ„Π΅Π΄Π΅ΠΏΠΈΠ½).

5) Амплитуда ΠœΠ‘Π“ Ρƒ ΠΌΠΎΠ»ΠΎΠ΄Ρ‹Ρ… ΠΌΡ‹ΡˆΠ΅ΠΉ Π»ΠΈΠ½ΠΈΠΈ DBA Π±Ρ‹Π»Π° Π²Ρ‹ΡˆΠ΅, Ρ‡Π΅ΠΌ Ρƒ C57BL6 ΠΌΡ‹ΡˆΠ΅ΠΉ, Ρ‡Ρ‚ΠΎ соотвСтствовало ΠΈΡ… ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡ‚ΠΈ ΠΊ ΠΎΠ±ΡƒΡ‡Π΅Π½ΠΈΡŽ. ΠŸΡ€Π΅Π΄ΠΏΠΎΠ»Π°Π³Π°Π΅Ρ‚ΡΡ Ρ‡Ρ‚ΠΎ ΠΌΠΎΠ»ΠΎΠ΄Ρ‹Π΅ DBA ΠΌΡ‹ΡˆΠΈ ΠΌΠΎΠ³ΡƒΡ‚ Π±Ρ‹Ρ‚ΡŒ ΠΈΡΠΏΠΎΠ»ΡŒΠ·ΠΎΠ²Π°Π½Π½Ρ‹ ΠΊΠ°ΠΊ модСль Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΡ ΠΎΠΏΡ€Π΅Π΄Π΅Π»Π΅Π½Π½Ρ‹Ρ… аспСктов обучСния ΠΈ ΠΏΠ°ΠΌΡΡ‚ΠΈ, связанных со ΡΡ‚Π°Ρ€Π΅Π½ΠΈΠ΅ΠΌ.

6) Π˜ΡΡ…ΠΎΠ΄Ρ ΠΈΠ· Π½Π°ΡˆΠΈΡ… Π΄Π°Π½Π½Ρ‹Ρ…, ΠœΠ‘Π“, ΠΈ Π”Π‘П ΠΏΡ€Π΅Π΄ΡΡ‚Π°Π²Π»ΡΡŽΡ‚ собой Π΄Π²Π° нСзависимых/ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°Ρ‚Π΅Π»ΡŒΠ½Ρ‹Ρ… Π½Π΅ΠΉΡ€ΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Ρ… ΠΌΠ΅Ρ…Π°Π½ΠΈΠ·ΠΌΠ° памяти, ΠΏΡ€ΠΈΡ‡Π΅ΠΌ ΡƒΠ²Π΅Π»ΠΈΡ‡Π΅Π½ΠΈΠ΅ Π½Π΅ΠΉΡ€ΠΎΠ½Π°Π»ΡŒΠ½ΠΎΠΉ возбудимости (ΠœΠ‘Π“) являСтся ΠΏΠ΅Ρ€Π²ΠΈΡ‡Π½Ρ‹ΠΌ ΠΏΠΎ ΠΎΡ‚Π½ΠΎΡˆΠ΅Π½ΠΈΡŽ ΠΊ Π΄Π»ΠΈΡ‚Π΅Π»ΡŒΠ½Ρ‹ΠΌ измСнСниям синаптичСской связи (Π”Π‘ΠŸ).

Π’Π°ΠΊ ΠΊΠ°ΠΊ старСниС ΠΌΠΎΠΆΠ΅Ρ‚ ΠΏΡ€ΠΈΠ²ΠΎΠ΄ΠΈΡ‚ΡŒ ΠΊ Ρ†Π΅Π»ΠΎΠΌΡƒ ряду Ρ„ΡƒΠ½ΠΊΡ†ΠΈΠΎΠ½Π°Π»ΡŒΠ½Ρ‹Ρ… Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠΉ, трСбуСтся Ρ€Π°Π·Π²ΠΈΡ‚ΠΈΠ΅ Π½ΠΎΠ²Ρ‹Ρ… ΠΌΠΎΠ΄Π΅Π»Π΅ΠΉ для изучСния Π½Π°Ρ€ΡƒΡˆΠ΅Π½ΠΈΠΉ обучСния ΠΈ ΠΏΠ°ΠΌΡΡ‚ΠΈ ΠΏΡ€ΠΈ старСнии.

ΠŸΠΎΠΊΠ°Π·Π°Ρ‚ΡŒ вСсь тСкст

Бписок Π»ΠΈΡ‚Π΅Ρ€Π°Ρ‚ΡƒΡ€Ρ‹

  1. Abdulla F.A., Abu-Bakra M.A.J., Calaminici M.R., Stephenson J.D., Sinden J.D. (1995) Importance of forebrain cholinergic and GABAergic systems to the aged-related deficits in water maze performance of rats. Neurobiol. Aging 16:41−52.
  2. W.C., Mason S.E., Demmer J., Williams J.M., Richardson C.L., Tate W.P., Lawlor P.A., Dragunow M. (1993) Correlation between immediate-early genes induction and the persistence of long-term potentiation. Neuroscience 56: 717−727.
  3. D.H., Meaney M.J. (1989) Temporally graded, age-related impairments in spatial memory in the rat. Neurobiol. Aging 10: 273−276.
  4. D.G., Witter M.P. (1989) The three-dimensional organization of the hippocampal formation: a review of anatomical data. J. Neurosci. 31(3): 571−591.
  5. P. (1963) Interhippocampal impulses. 1. Basal dendritic activation of CA1 neurons. Acta physiol. Scand. 48: 178−208.
  6. P., Eccles J.C., Loyning Y. (1963) Recurrent inhibition in the hippocampus with identification of the inhibitory cell and its synapses. Nature 198: 540−542.
  7. P., Eccles J.C., Loyning Y. (1964a) Location of postsynaptic inhibitory synapses on hippocampal pyramids. J. Neurophysiol. 27: 592−607.
  8. P., Eccles J.C., Loyning Y. (1964b) Pathway of postsynaptic inhibition in the hippocampus. J. Neurophysiol. 27: 608−619.
  9. P., Holmquist B., Voorholve P.E. (1968) Excitatory synapses on hippocampal apical dendrites activated by enthorinal stimulation. Acta physiol. Scand. 66(4): 461−473.
  10. P., Bliss T.V., Skrede K.K. (1971) Lamellar organization of hippocampal excitatory pathways. Exp. Brain Res. 13: 222−238.
  11. R., Nicoll R.A. (1987) Pharmacologically distinct actions of serotonin on single pyramidal neurones of the rat hippocampus recorded in vitro. J Physiol. 394: 99 124.
  12. Anikszejn L., Ben-Ari Y. (1991) Novel form of long-term potentiation produced by K+ channel blocker in hippocampus. Nature. 341:67−69.
  13. A., Singer W. (1993) Long-term depression of excitatory synaptic transmission and its relationship to long-term potentiation. Trends Neurosci. 16:480−487.
  14. J., Riekkinen M., Riekkinen P. Jr. (1998) Tetrahydroaminoacridine and D-cycloserine stimulate acquisition of water maze spatial navigation in aged rats. Eur. J. Pharmacol. 342: 15−20.
  15. P.V., Gerber M., Joseph J.A., Crews F.T. (1998) Uncoupling of muscarinic cholinergic phosphoinositide signals in senescent cerebral cortical and hippocampal membranes. Neurochem Int. 32(1): 107−15.
  16. C.A. (1979) Memory deficits associated with senescence: a neurophysiological and behavioral study in the rat. J. Comp. Physiol. Psychol. 93:74−104.
  17. C.A. (1988) Aging and the physiology of spatial memory. Neurobiol. Aging. 9: 563−568.
  18. C.A. (1990) Animal models of age-related cognitive decline. In: Boiler F., Grafman J. (Eds.), Handbook of Neuropsychology. Elsevier, Amsterdam, pp. 169−196.
  19. C.A. (1999) Do synaptic markers provide a window on synaptic effectiveness in the aged hippocampus? Neurobiol. Aging 20: 349−351.
  20. C.A. (1994) Normal aging: regionally specific changes in hippocampal synaptic transmission. Trends Neurosci. 17: 13−18.
  21. Barnes C.A., McNaughton B.L. (1979) Neurophysiological comparison of dendritic cable properties in adolescent, middle-aged, and senescent rats. Exp. Brain Res. 5:195 206.
  22. Barnes C.A., McNaughton B.L. (1980a) Physiological compensation for loss of afferent synapses in rat hippocampus granule cells during senescence. J. Physiol. (London) 309: 473−485.
  23. Barnes C.A., McNaughton B.L. (1980b) Spatial memory and hippocampal synaptic plasticity in middle-aged and senescent rats. In: Stein, D. (Ed.), Psychobiology of Aging: Problems and Perspectives. Elsevier, New York, pp. 253−272.
  24. Barnes C.A., McNaughton B.L. (1985) An age comparison of the rates of acquisition and forgetting of spatial information in relation to long-term enhancement of hippocampal synapses. Behav. Neurosci. 99:1040−1048.
  25. C.A., Nadel L., Honig W.K. (1980) Spatial memory deficit in senescent rats. Can. J. Psychol. 34: 29−39.
  26. Barnes C.A., McNaughton B.L., O’Keefe J. (1983) Loss of place specificity in hippocampal complex spike cells of senescent rat. Neurobiol. Aging 4:113−119.
  27. Barnes C. A, Rao G., Foster T.C., McNaughton B.L. (1992) Region-specific age effects on AMPA sensitivity: electrophysiological evidence for loss of synaptic contacts in hippocampal field CA1. Hippocampus. 2(4): 457−68.
  28. Barnes C.A., Rao G., Shen J. (1997a) Age-related decrease in the N-methyl-D-aspartateR-mediated excitatory postsynaptic potential in hippocampal region CA1. Neurobiol. Aging 18:445−452.
  29. Barnes C.A., Rao G., McNaugthon B.L. (1996) Functional integrity of NMDA-dependent LTP induction mechanisms across the lifespan of F344 rats. Learn. Mem. 3: 124−137.
  30. Barnes C.A., Suster M.S., Shen J., McNaughton B.L. (1997b) Multistability of cognitive maps in the hippocampus of old rats. Nature 388: 272−275.
  31. Barnes C.A., Meltzer J., Houston F., Orr G., McGann K., Wenk G.L. (2000a) Chronic treatment of old rats with donepezil or galantamine: effects on memory, hippocampal plasticity and nicotinic receptors. Neuroscience 99: 17−23.
  32. Barnes C.A., Rao G., Houston F.P. (2000b) LTP induction threshold change in old rats at the perforant path-granule cell synapse. Neurobiol. Aging 21: 613−620.
  33. Barnes C.A., Rao G., Orr G. (2000c) Age-related decrease in the Schaffer collateral-evoked EPSP in awake, freely behaving rats. Neural Plast. 7: 167−178.
  34. A., Muller D., Derkach V., Griffith L.C., Soderling T.R. (1997) Regulatory phosphorylation of AMPA-type glutamate receptors by CaMKlI during long-term potentiation. Science 276: 2043−2045.
  35. Baxter M.G., Lanthorn T.H., Frick K.M., Golski S., Wan R.Q., Olton D.S. (1994) D-Cycloserine, a novel cognitive enhancer, improves spatial memory in aged rats. Neurobiol. Aging 15: 207−213.
  36. M.F., Abraham W.C. (1996) Long-term depression in hippocampus. Annu Rev Neurosci. 19: 437−62. Review.
  37. Belcadi-Abassi W., Destrade C. (1995) Post-test apamin injection suppresses a Kamin-like effect following a learning session in mice. NeuroReport 6: 437−462.
  38. T.A., Luthi A., Isaac J.T., Collingridge G.L. (1998) Modulation of AMPA receptor unitary conduction by synaptic activity. Nature 393: 793−797.
  39. L.S., Prince D.A. (1982) Cholinergic excitation of mammalian hippocampal pyramidal cells. Brain Res. 249(2): 315−31.
  40. Bickford-Wimcr P.C., Miller J. A, Freedman R., Rose G.M. (1988) Age-related reduction in responses of rat hippocampal neurons to locally applied monoamines. Neurobiol Aging. 9(2): 173−9.
  41. E.L., Cooper L.N., Munro P.W. (1982) Theory for the development of neuron selectivity: orientation specificity and binocular interaction in visual cortex.1. J Neurosci. 2(1): 32−48.
  42. Bliss T.V.P., Gardner-Medwin A.R. (1973) Long-lasting potentiation of synaptic transmission in the dentate area of the unanaesthetised rabbit following stimulation of the perforant path. J. Physiol. (London) 232: 357−374.
  43. Bliss T.V.P. Lomo T. (1973) Long-lasting potentiation of synaptic transmission in the dentate area of the anaesthetised rabbit following stimulus of the perforant path. J. Physiol. (London) 232: 331−356.
  44. Bliss T.V.P, Collingridge G.L. (1993) A synaptic model of memory: long-term potentiation in the hippocampus. Nature 361: 31−39.
  45. Bliss T.V.P. (2003) A journey from neocortex to hippocampus. Philos. Trans. R. Soc. Lond B Biol. Sci. 358: 621−623.
  46. R.D., Wong T., Nouranifar R., Iyengar R., Landau E.M. (1995) Postsynaptic cAMP pathway gates early LTP in hippocampal CA1 region. Neuron. 15: 1403−1414.
  47. R.D., Connor J.H., Brown G.P., Wong T., Shenokilar S., Iyengar R., Landau E.M. (1998) Gating of CaMKII by cAMP-regulated protein phosphatase activity during LTP. Science. 280: 1940−1942.
  48. A. (1996) Acetylcholine: a neurotransmitter for learning and memory? Brain Res. Rev. 21:285−300.
  49. S.E., Fletcher S., Loane D.J., Marrion N.V. (2001) Somatic colocalization of rat SKI and D class (Ca (v)1.2) L-type calcium channels in rat CA1 hippocampal pyramidal neurons. J Neurosci. 21(20): RC175.
  50. Y., Solntseva E.L., Skrebitsky V.G. (2002) Long-term potentiation of the glutamate-activated inward current induced by 8-Br-cGMP in nerve cell.1. Dokl Biol Sci. 384:191−4.
  51. R.D., Gallagher M. (1993) A longitudinal study of reaction time performance in Long-Evans rats. Neurobiol. Aging 14: 57−64.
  52. S.R. (1955) Studies on the cerebral cortex (limbic structures). London.
  53. Campbell L. W, Hao S.Y., Thibault O., Blalock E.M., Landfield P.W. (1996) Aging changes in voltage-gated calcium currents in hippocampal CA1 neurons. J Neurosci. 16(19):6286−95.
  54. I., Teyler T.J. (1996) Two forms of long-term potentiation in area CA1 activate different signal transduction pathways. J. Physiol. (London) 76:3038−3047.
  55. Charpak S., Gahwiler B.H., Do K.Q., Knopfel T. (1990) Potassium conductances in hippocampal neurons blocked by excitatory amino-acid transmitters. Nature. 47(6295): 765−7.
  56. C., Tonegava S. (1997) Molecular genetic analysis of synaptic plasticity, activity-dependent neural development, learning and memory in mammalian brain. Annu. Rev. Neurosci. 20:157−184.
  57. D.A., Browning M.D. (2001) Deficits in the expression of the NR2B subunit in the hippocampus of aged Fisher 344 rats. Neurobiol Aging. 22(1): 165−8.
  58. R.K., Tavalin S.J., Marrion N.V. (1997) Beta-adrenergic stimulation selectively inhibits long-lasting L-type calcium channel facilitation in hippocampal pyramidal neurons. J Neurosci. 17(17): 6493−503.
  59. A.S., Raymond C.R., Abraham W.C. (1998) Priming of long-term potentiation induced by activation of metabotropic glutamate receptors coupled to phospholipase C. Hippocampus. 8(2): 160−70.
  60. A.E., Nicoll RA. (1983) Acetylcholine mediates a slow synaptic potential in hippocampal pyramidal cells. Science. 221(4617): 1299−301.
  61. A., Halliwell J.V. (1987) Differential modulation of three separate K-conductances in hippocampal CA1 neurons by serotonin. Nature. 328(6125): 73−7.
  62. Collingridge G.L., Lester R.A.J. (1989) Excitatory amino acide receptors in the vertebrate central neurvous system. Pharmacol. Rev. 40: 143−210.
  63. Collingridge G.L., Kehl, S.J., McLennan H. (1983a) Excitatory amino acids in synaptic transmission in the Schaffer collateral-commissural pathway of the rat hippocampus. J. Physiol. (London) 334: 33−46.
  64. P.J., Wetsel W.C., Gallagher M. (1997) Spatial memory is related to hippocampal subcellular concentrations of calcium-dependent protein kinase C isoforms in young and aged rats. Proc. Natl. Acad. Sci. USA 94: 14 195−14 199.
  65. C.M., Teyler T.J. (1996a) Long-term potentiation induces synaptic plasticity at nontetanized adjacent synapses. Learn. Mem. 3:106−114.
  66. C.M., Teyler T.J. (1996b) Protein kinase and phosphatase activity regulate the form of synaptic plasticity expressed. Synapse 24(2):97-l 03.
  67. R.H., Turrigiano G.G. (2004) Long-term potentiation of intrinsic excitability in LV visual cortical neurons. J Neurophysiol 92:341−348.
  68. G., Debanne D. (2003) Long-term plasticity of intrinsic excitability: learning rules and mechanisms. Learn. Mem. 10: 456−465.
  69. Decker M.W., McGaugh J.L. (1991) The role of interactions between the cholinergic system and other neuromodulatory systems in learning and memory. Synapse. 7(2): 151−68. Review.
  70. De Jong G.I., Buwalda B., Schuurman T., Luiten P.G. (1992) Synaptic plasticity in the dentate gyrus of aged rats is altered after chronic nimodipine application. Brain Res 596:345−348.
  71. O., Bizot J.C. (1997) Effect of apamin, a selective blocker of Ca2±activated K+ channel, on habituation and passive-avoidance responses in rats. Neurosci. Lett. 222: 57−60.
  72. O., Bizot J.C., Goyffron M. (1997) Apamin improves learning in an object recognition task in rats. Neurosci. Lett. 222: 159−162.
  73. D.L., Turner D.A., Watters C.L. (1991) Spatial performance correlates with in vitro potentiation in young and aged Fischer 344 rats. Brain Res. 554:1−9
  74. D.L., Bradley D.A., Turner D.A., (1993) Age-related alternations in potentiation in the CA1 region in F344 rats. Neurobiol. Aging 14: 249−258.
  75. R.A., Straube K.T., Disterhoft J.F. (1989) Nimodipine facilitates associative learning in aging rabbits. Science 243:809−811.
  76. Diana G., Domenici M.R., Loizzo A., Scotti de Carolis A., Sagratella S. (1994a) Age and strain differences in rat place cell learning and hippocampal dentate gyrus frequency potentiation. Neurosci. Lett. 171:113−116.
  77. Diana G., Scotti de Carolis A., Frank C., Domenici M.R., Sagratella S (1994b) Selective reduction of hippocampal dentate frequency potentiation in aged rats with impaired place learning. Brain Res. Bull. 35:107−111.
  78. J.F., Coulter D.A., Alkon D.L. (1986) Conditioning-specific membrane changes of rabbit hippocampal neurons measured in vitro. Proc Natl Acad Sci USA. 83(8): 2733−7.
  79. Disterhoft J.F., Moyer J.R. Jr, Thompson L.T. (1994) The calcium rationale in aging and Alzheimer’s disease. Evidence from an animal model of normal aging. Ann N Y Acad Sci. 15- 747: 382−406. Review.
  80. Disterhoft J. F, Thompson L.T., Moyer J.R. Jr, Mogul D.J. (1996) Calcium-dependent afterhyperpolarization and learning in young and aging hippocampus. Life Sci. 59(5−6): 413−20. Review.
  81. Disterhoft J.F., Wu W.W., Ohno M. (2004) Biophysical alterations of hippocampal pyramidal neurons in learning, aging and Alzheimer disease. Ageing Res. Rev. 3: 383 406.
  82. R.M., Goddard G.V. (1975) Long-term potentiation of the perforant path-granule cell synapse in the rat hippocampus. Brain. Res. 86: 205−215.
  83. S.M., Bear M.F. (1993) Bidirectional long-term modification of synaptic effectiveness in the adult and immature hippocampus. J Neurosci. 13(7): 2910−8.
  84. S.B., Evendcn J.L., Iversen S.D. (1988) Delay-dependent short-term memory deficits in aged rats. Psychopharmacology 96: 174−180.
  85. S.B., Martel F.L., Iversen S.D. (1990) Proactive interference effects on short-term memory in rats: II. Effects in young and aged rats. Behav. Neurosci. 104: 666 670.
  86. Eckles-Smith K., Clayton D., Bickford P., Browning M.D. (2000) Caloric restriction prevents age-related deficits in LTP and in NMDA receptor expression. Brain Res Mol Brain Res. 78(1−2): 154−62.
  87. Faber E.S., Sah P. (2003) Ca2±activated K+ (BK) channel inactivation contributes to spike broadening during repetitive firing in the rat lateral amygdala. J Physiol 552:483 497.
  88. N.B., Sergeeva O.A., Skrebitsky V.G. (1993) Priming stimulation facilitates Hebb-type plasticity in the Schaffer collateral-commissural pathways of the mouse hippocampus. Exp Brain Res. 94(2): 270−2.
  89. A. Boddeke H., Mueller D. (1992) Enhancement of AMPA-mediated synaptic transmission by protein phosphatase inhibitor calyculin A in rat hippocampal slices. Eur. J. Neurosci. 2: 1035−1041.
  90. W., Chen K.S., Gage F.H., Bjorklund A. (1991) Progressive decline in spatial learning and integrity of forebrain cholinergic neurons in rats during aging. Neurobiol. Aging 13: 9−23.
  91. K.M., Mogul D.J. (1997) Adenosine A3 receptors potentiate hippocampal calcium current by a PKA-dependent/PKC-independent pathway. Neuropharmacology. 36(3): 353−62.
  92. D.E., Wehner J.M. (1993) Physical activity enhances spatial learning performance with an associated alteration in hippocampal protein kinase Π‘ activity in C57BL/6 and DBA/2 mice. Brain Res. 619(1−2): 111−9.
  93. D.E., Clark V.J., Paylor R., Wehner J.M. (1995) Enhancement of hippocampally-mediated learning and protein kinase C activity by oxiracetam in learning-impaired DBA/2 mice. Brain Res. 672(1−2): 170−6.
  94. Foster T.C., Barnes C.A., Rao G., McNaughton B.L. (1991) Increase in perforant path quantal size in aged F-344 rats. Neurobiol. Aging. 12: 441−448.
  95. T.C., Norris C.M. (1997) Age-associated changes in Ca(2+)-dependent processes: relation to hippocampal synaptic plasticity. Hippocampus 7:602−612.
  96. T.C., Norris C.M. (1998) On slices, synaptosomes and dissociated neurones to study in vitro ageing physiology. Trends Neurosci. 21(7): 286−7.
  97. T.C., Fugger H.N., Cunningham S.G. (1999) Experience-dependent enhancement of perforant path synaptic transmission: relation to hippocampal function. Submitted.
  98. T.C., Sharrow K.M., Masse J.R., Norris C.M., Kumar A. (2001) Calcineurin links Ca2+ dysregulation with brain aging. J Neurosci. 21(11):4066−73.
  99. T.C., Kumar A. (2002) Calcium dysregulation in the aging brain. Neuroscientist 8:297−301.
  100. Frankland P.W., O’Brien C., Ohno M., Kirkwood A., Silva A.J. (2001) Alpha-CaMKII-dependent plasticity in the cortex is required for permanent memory. Nature. 411:309−313.
  101. K.M., Baxter M.G., Markowska A.L., Olton D.S., Price D.L. (1995) Age-related spatial reference and working memory deficits assessed in the water maze. Neurobiol. Aging 16:149−160.
  102. Frick K.M., Stillner E.T., Berger-Sweeney J. (2000) Mice are not little rats: species differences in a one-day water maze task. Neuroreport. 11(16): 3461−5.
  103. A., Johnston D. (2005) Plasticity of dendritic excitability. J. Neurobiol. 64: 100−115.
  104. H.N., Lichtenvoort J.M., Foster T.C. (1997) Enthorhinal cortex lesions as a model of age-related changes in hippocampal function. Psychobiology 25: 277−285.
  105. Gage F.H., Bjorklund A., Stenevi U., Dunnett S.B., Kelly P.A.T. (1984a) Intrahippocampal spatial grafts ameliorate learning impairments in aged rats. Science. 225:533−536.
  106. F.H., Dunnett S.B., Bjorklund A. (1984b) Spatial learning and motor deficits in aged rats. Neurobiol. Aging. 5:43−48.
  107. Gage F.H., Kelly P.A.T., Bjorklund A. (1984c) Regional changes in brain glucose metabolism reflect cognitive impairments in aged rats. J. Neurosci. 4:2856−2865.
  108. F.H., Dunnett S.B., Bjorklund A. (1989) Age-related impairments in spatial memory are independent of those in sensory-motor skills. Neurobiol. Aging. 10:347−352.
  109. M., Colombo P.J. (1995) Aging: the cholinergic hypothesis of cognitive decline. Curr. Opin. Neurobiol. 5:161−168.
  110. M., Pelleymounter M.A. (1988) Spatial learning deficits in old rats: a model for memory decline in the aged. Neurobiol. Aging. 9:363−369.
  111. M., Bostock E., King R. (1985) Effects of opiate antagonists on spatial memory in young and aged rats. Behav. Neural Biol. 44:374−385.
  112. Gallagher M., Burwell R.D., Kodsi M.H., McKinney M., Southerland S., Vella-Rountree L., Lewis M.H. (1990) Markers for biogenic amines in the aged rat brain: relationship to decline in spatial learning ability. Neurobiol. Aging. 11:507−514.
  113. M., Burwell R.D., Burchinal M. (1993) Severity of spatial learning impairment in aging: development of a learning index for performance in Morris Water Maze. Behav. Neurosci. 107(4): 618−626.
  114. Y. (1979) Loss of axosomatic synapses in the dentate gyrus of aged rats. Brain Res. 168:485−492.
  115. Geinisman Y., de Toledo-Morrell L., Morrell F. (1986) Aged rats need a preserved complement of perforated axospinous synapses per hippocampal neuron to maintain good spatial memory. Brain Res. 398(2): 266−75.
  116. Geinisman Y., de Toledo-Morrell L., Morrell F., Heller R.E. (1995) Hippocampal markers of age-related memory dysfunction: behavioral, electrophysiological and morphological perspectives. Prog. Neurobiol. 45: 223−252.
  117. U., Gahwiler B.H. (1994) Modulation of potassium conductances by metabotropic glutamate receptors in the hippocampus. Ren Physiol Biochem. (3−4): 12 931.
  118. Giese K. P, Fedorov N.B., Filipkowsli R.K., Silva A.J. (1998) Autophosphorylation at Thr286 of the alpha-calcium-calmodulin kinase II in LTP and learning. Science 279: 870−873.
  119. Giese K. P, Peters M., Vernon J. (2001) Modulation of excitability as a learning and memory mechanism: a molecular genetic perspective. Physiol. Beh. 73: 803−810.
  120. Gold P.E., McGaugh J.L., Hankins L.L., Rose R.P., Vasques B.J. (1981) Age-dependent changes in retention in rats. Exp. Aging Res. 8: 53−58.
  121. J.D. (1960) The hippocampus. In: Handbook of physiology1.Neurophysiology. Ed. Fiels, Magoun and Hall. 2. Baltimore, Williams and Wilkins, 1373−1389.
  122. J.D., Maxwell D.S. (1961) Hippocampal electrical activity: 1. Morphological aspects. EEG clin. Neurophysiol., 13: 837−846.
  123. Grover L. M (1998) Evidence for postsynaptic induction and expression ofNMDA receptor independent LTP. J Neurophysiol. 79(3): 1167−82.
  124. L.M., Teyler T.J. (1990) Two components of long-term potentiation induced by different patterns of afferent activation. Nature 347:447−479.
  125. L.M., Teyler T.J. (1992) N-Methyl-D-aspartate receptor-independent long-term potentiation in area CA1 of rat hippocampus: input-specific induction and preclusion in a non-tetanized pathway. Neuroscience, 49:7−11.
  126. L.M., Teyler T.J. (1994) Activation ofNMDA receptors in hippocampal area CA1 by low and high frequency orthodromic stimulation and their contribution to induction of long-term potentiation. Synapse. 16:66−75.
  127. B., Wigstrom H. (1981) Shape of frequency-current curves in CAI pyramidal cells in the hippocampus. Brain Res. 223(2): 417−21.
  128. B., Wigstrom W.C. (1988) Physiological mechanisms underlying long-term potentiation. Trends Neurosci. 11: 156−162.
  129. H.L., Konnerth A. (1983) Histamine and noradrenaline decrease calcium-activated potassium conductance in hippocampal pyramidal cells. Nature. 302(5907): 432−4.
  130. H.L., Greene R.W. (1984) Adenosine enhances afterhyperpolarization and accommodation in hippocampal pyramidal cells. Pflugers Arch. 402(3): 244−7.
  131. Hayashi Y., Shi S.H., Esteban J.A. Piccini A., Poncer J.C., Malinow R. (2000) Driving AMPA receptors into synapses by LTP and CaMKII: requirement for GluRl and PDZ domain interaction. Science 287: 2262−2267.
  132. D.O. (1949) The organization of behavior. Wiley, New York.
  133. J.W., Yokoyama C.T., Breeze L.J., Chavkin C., Catterall W.A. (1995) Phosphorylation of presynaptic and postsynaptic calcium channels by cAMP-dependent protein kinase in hippocampal neurons. EMBO J. 14(13):3036−44.
  134. Heyen A.J., Quinlan E.M. Bae D.C., Bear M.F. (2000) Bidirectional, activity-dependent regulation of glutamate receptors in the adult hippocampus in vivo. Neuron 28: 527−536.
  135. G.E., Anderson J.A. (1981) Parallel Models of Associative Memory. Lawrence Erlbaum Associates, Hillsdale.
  136. Hjorth-Simonsen A.(1972) Projection of the lateral part of the enthorinal area to the hippocampus and fascia dentate. J. Comp. Neurol., 146(2): 219−232.
  137. Hjorth-Simonsen A. (1976) Laminar distribution and topical organization of intrinsic connections in the hippocampal region. Exp. Brain Res., Suppl. 1: 171−176.
  138. J.J. (1982) Neural networks and physical systems with emergent collective computational abilities. Proc. Natl. Acad. Sci. USA 79: 2554−2558.
  139. J.R., Prince D.A. (1980) A calcium-activated hyperpolarization follows repetitive firing in hippocampal neurons. J Neurophysiol. 43(2): 409−19.
  140. S., Sacktor T.C. (1996) Bidirectional regulation of protein kinase M zeta in the maintenance of long-term potentiation and long-term depression. J Neurosci. 16(17):5324−33.
  141. P.T., Lisman J.E. (1995) Bidirectional synaptic plasticity induced by a single burst during cholinergic theta oscillation in CA1 in vitro. Neuron. 15(5): 1053−63.
  142. Y.Y., Kandell E.R. (1994) Recruitment of long-lasting and protein kinase-A-dependent long-term potentiation in the CA1 region of hippocampus requires repeated tetanization. Learn Mem. 1: 74−82.
  143. Isaac J.T.R., Nicoll R.A., Malenka R.C. (1995) Evidence for silent synapses: implication for the expression of LTP. Neuron 15:427−434.
  144. Ingram D.K., Spangler E.L., Iijima S., Kuo H., Bresnahan E.L., Greig N.H., London E.D. (1994) New pharmacological strategies for cognitive enhancement using a rat model of age-related memory impairment. Ann. N.Y. Acad. Sci. 717: 16−32.
  145. D., Kornisiuk E., Izquierdo I. (1997) Cholinergic neurotransmission and synaptic plasticity concerning memory processing. Neurochemistry Res. 22:507−515.
  146. M.W., Peckham H.M., Errington M.L., Bliss T.V., Routtenberg A. (2001) Synaptic plasticity in the hippocampus of awake C57BL/6 and DBA/2 mice: interstrain difference and parallels with behavior. Hippocampus. 11(4): 391−396.
  147. D., Williams S., Jaffe D., Gray R. (1992) NMDA-receptor independent long-term potentiation. Annu. Rev. Physiol. 54: 489−505.
  148. H., Izumi Y., Zorumski C.F. (1997) Removal of extracellular calcium after conditioning stimulation disrupts long-term potentiation in the CA1 region of rat hippocampal slices. Neuroscience. 76(4): 1113−9.
  149. Z.S. (1984) Towards theories of brain aging. In: Kay, D., Burrows, G.D. (Eds.), Handbook of Studies on Psychiatry and Old Age. Elsevier, Amsterdam, pp. 7−30.
  150. Z.S. (1989) The role of calcium regulation in brain aging: reexamination of a hypothesis. Aging (Milano). 1(1): 17−34. Review. Erratum in: Aging (Milano) 1(2):II.
  151. Z.S. (1994) Calcium hypothesis of Alzheimer’s disease and brain aging. Ann N Y Acad Sci. 747: 1−11. Review.
  152. J. A., Marshall G.R., Woodruff G.N. (1986) Quantitative evaluation of the potencies of GABA-receptor agonists and antagonists using the rat hippocampal slice preparation. Br. J. Pharmac., 87, 677−684.
  153. T. (1978) Associative Memory: A System-Theoretic Approach. SpringerVerlag, New York.
  154. M., Disterhoft J.F. (1994) Relation of nimodipine dose and serum concentration to learning enhancement in aging rabbits. Exp Neurol. 127(1): 159−66.
  155. M., Pedarzani P. (2000) A protein phosphatase is involved in the cholinergic suppression of the Ca2±activated K+ current sI(AHP) in hippocampal pyramidal neurons. Neuropharmacology 39: 1274−1293.
  156. Kronforst-Collins M.A., Moriearty P.L., Schmidt B., Disterhoft J.F. (1997) Metrifonate improves associative learning and retention in aging rabbits. Behav Neurosci. 111(5): 1031−40.
  157. B., Adams P.R. (1986) Calcium-dependent current generating the afterhyperpolarization of hippocampal neurons. J Neurophysiol. 55(6): 1268−82.
  158. P.W. (1988) Hippocampal neurobiological mechanisms of age-related memory dysfunction. Neurobiol. Aging 9: 571−579.
  159. P.W. (1994) Increased hippocampal Ca2+ channel activity in brain aging and dementia. Hormonal and pharmacologic modulation. Ann N Y Acad Sci. 747: 351 -64. Review.
  160. P.W. (1996) Aging-related increase in hippocampal calcium channels. Life Sci 59:399−404.
  161. P.W., Lynch G. (1977) Impaired monosynaptic potentiation in vitro hippocampal slices from of aged memory-deficient rats. J. Gerontol. 150: 85−101.
  162. P.W., Pitler T.A. (1984) Prolonged Ca2±dependent afterhyperpolarizations in hippocampal neurons of aged rats. Science. 226(4678): 108 992.
  163. Landfield P.W., McGaugh J.L., Lynch G. (1978) Imapaired synaptic potentiation process in the hippocampus of aged, memory-deficient rats. Brain. Res. 32: 523−533.
  164. Lee H.K., Barbaroise M., Kameyama K., Bear M.F., Huganir R.L. (2000) Regulation of distinct AMPA receptor phosphorylation sites during bidirectional synaptic plasticity. Nature 405: 955−959.
  165. W.B., Steward O. (1979) Synapses as associative memory elements in the hippocampal formation. Brain Res. 175: 233−245.
  166. Li C., Lu J., Wu C., Duan S., Poo M. (2004a) Bidirectional modification of presynaptic neuronal excitability accompanying spike timing-dependent synaptic plasticity. Neuron 41:257−268.
  167. Li C., Lu J., Wu C., Duan S., Poo M. (2004b) Bidirectional modification of presynaptic neuronal excitability accompanying spike timing-dependent synaptic plasticity. Neuron 41:257−268.
  168. D., Hessler N.A., Malinow R. (1995) Activation of postsynaptically silent synapses during LTP in CA1 region of hippocampal slice. Nature 375: 400−404.
  169. D.L., Mody I. (1994) Regulation ofNMDA channel function by endogenous Ca2±dependent phosphatase. Nature 369: 235−239.
  170. D.J., Connors J.A. (1995) Ling-term synaptic depression. Annu. Rev. Neurosci. 18:319−358.
  171. M.D., Schallert T. (1988) Aging and atropine effects on spatial navigation in the Morris water task. Behav. Neurosci. 102: 621−634.
  172. J. (1989) A mechanism for the Hebb and the anti-Hebb processes underlying learning and memory. Proc Natl Acad Sci USA. 86(23):9574−8.
  173. J., Schulman H., Cline H. (2002) The molecular basis of CaMKII function in synaptic and behavioral memory. Nat. Rev. Neurosci. 3: 175−190.
  174. P.M., Hjelmstad G.O., Mukheiji S., Soderling T.R., Malenka R.C., Nicoll R.A. (1995) Calcium/calmodulin-dependent kinase II and long-term potentiation enhance synaptic transmission by the same mechanism. Proc Natl Acad Sci USA. 92(24): 111 759.
  175. Lorente de No (1934) Junctional intercellular communication: the cell-to-cell membrane channel. Physiol. Rev., 61:829−913.
  176. D.M., Routtenberg A. (1987) Protein kinase C stimulators produce a synapse specific increase in the endurance of long-term potentiation. Brain Res. 436: 177−183.
  177. C., Frerking M. (2001) Restless AMPA reveptors: implications for synaptic transmission and plasticity. Trends Neurosci. 24: 665−670.
  178. G., Baudry M. (1984) The biochemistry of memory: a new and specific hypothesis. Science 224: 1057−1063.
  179. G., Voss K.L. (1994) Membraine arachidonic acid concentration correlates with age and induction of long-term potentiation in the dentate gyrus of the rat. Eur. J. Neurosci. 6: 1008−1014.
  180. D.V., Nicoll R.A. (1982) Noradrenaline blocks accommodation of pyramidal cell discharge in the hippocampus. Nature. 299(5884): 636−8.
  181. J.C. (2002) Synaptic and non-synaptic mechanisms for plasticity and learning. In: Proceedings of the Winter Conference on Neural Plasticity. Moorea, French Polynesia.
  182. R.C., Kauer J.A., Perkel D.J., Mauk M.D., Kelly P.T., Nicoll R.A., Waxham M.N. (1989) An essential role for postsynaptic calmodulin and protein kinase activity in long-term potentiation. Nature. 340(6234): 554−7.
  183. R.C., Nicoll R.A. (1999) Long-term potentiation a decade of progress? Science 285: 1870−1874.
  184. R., Madison D.V., Tsien R.W. (1988) Persistent protein kinase activity underlying long-term potentiation. Nature. 335 (6193): 820−4.
  185. R., Schulman H., Tsien R.W. (1989) Inhibition of postsynaptic PKC or CaMKII blocks induction but not expression of LTP. Science 245: 862−865.
  186. R., Malenka R.C. (2002) AMPA receptor trafficking and synaptic plasticity. Annu. Rev. Neurosci. 25: 103−126.
  187. Marby T.R., McCarty R., Gold P.E., Foster T.C. (1996) Age and stress-history effects on spatial performance in a swim task in Fischer 344 rats. Neurobiol. Learn. Mem. 66: 1−10.
  188. A.L., Stone W.S., Ingram D.K., Reynolds J., Gold P.E., Conti L.H., Pontecorvo M.J., Wenk J.L., Olton D.S. (1989) Individual differences in aging: behavioral and neurobiological correlates. Neurobiol. Aging 10: 31−43.
  189. D. (1971) Simple memory: a theory for archicortex. Philos. Trans. R. Soc. B: Biol. Sci. 262:23−81.
  190. N.V., Tavalin S.J. (1998) Selective activation of Ca2±activated K+ channels by co-localized Ca2+ channels in hippocampal neurons. Nature. 395(6705): 900−5.
  191. Mayford M., Wang J., Kandel E.R., O’Dell T.J. (1995) CaMKII regulates the frequency-response function of hippocampal synapses for the production of both LTD and LTP. Cell. 81(6): 891−904.
  192. J.L., Rigter H. (1983) Assessment of retention capacities in old rats. Behav. Neural Biol. 39: 181−191.
  193. McCann S.M. (1997) The nitric oxide hypothesis of brain aging. Exp Gerontol. 32(4−5): 431−40. Review
  194. McClelland J.L., Rumelhart D.E. (1986) Parallel Distributed Processing: Explorations in the Microstructure of Cognition. MIT Press, Cambridge.
  195. McGahon B.M., Clements M.P., Lynch G. (1997) The ability of aged rats to sustain long-term potentiation is restored when the age-related decrease in membrane arachidonic acid concentration is reversed. Neuroscience. 81: 9−16.
  196. McGaugh J. L, (1989) Involvement of hormonal and neuromodulatory systems in the regulation of memory storage. Annu. Rev. Neurosci. 12:255−287.
  197. H. (1961) Techniques in tissue metabolism. 5. Chopping and slicing tissue samples. Biochem. J. 27: 213−218.
  198. H., Batchelard H.S. (1976) Biochemistry and the central nervous system. 4th edition. Churchill Livingston, London.
  199. McNaughton B.L., Barnes C.A. (1977) Physiological identification and analysis of DG cell responses to stimulation of the medial and lateral perforant pathways in the rat. Comparativ. Neur. 175(4): 439−451.
  200. McNaughton B.L., Douglas R.M., Goddard G.V. (1978) Synaptic enhancement in fascia dentate: cooperativity among coactive afferents. Brain Res. 157: 277−293.
  201. McNaughton B.L., Morris R.V.M. (1987) Hippocampal synaptic enhancement and information storage within a distributed memory system. Trends Neurosci. 10: 408−415.
  202. Monagle-Strucko K, Fanelli RJ (1993) Enhanced acquisition of reversal training in a spatial learning task in rats treated with chronic nimodipine. Pharmacol Biochem Behav 44:827−835.
  203. C.I., Browning M.D., Rose G.M. (1993) Hippocampal plasticity induced by prime burst, but not long-term potentiation, stimulation is impaired in area CA1 of aged Fischer 344 rats. Hippocampus 3: 57−66.
  204. S.L., Teyler T.J. (1999) Epileptic-like activity induces multiple forms of plasticity in hippocampal area CA1. Brain Res. 917(1): 90−6.
  205. S.L., Teyler T.J. (2001 a) Electrical stimuli patterned after the theta-rhythm induce multiple forms of LTP. J Neurophysiol. 86(3): 1289−96.
  206. S.L., Teyler T.J. (2001b) VDCCs and NMDARs underlie two forms of LTP in CA1 hippocampus in vivo. J Neurophysiol. 82(2): 736−40.
  207. S.L., Coussens C.M., Teyler T.J. (2001) Depotentiation of vdccLTP requires NMDAR activation. Neurobiol Learn Mem. 76(3): 229−38.
  208. Morris R.G.M. (1981) Spatial localization does not require the presence of local cues. Learn. Motiv. 12:239−261.
  209. Morris R.G.M. (1989) Synaptic plasticity and learning: selective impairment of learning in rats and blockade of long-term potentiation in vivo by the 7V-methyl-D-aspartate receptor antagonist D-AP5. J. Neurosci. 2: 1016−1028.
  210. R.G., Frey U. (1997) Hippocampal synaptic plasticity: role in spatial learning or the automatic recording of attended experience? Philos. Trans. R. Soc. London, Ser. B Biol. Sci. 352: 1489−1503.
  211. Moyer J.R. Jr, Thompson L.T., Black J.P., Disterhoft J.F. (1992) Nimodipine increases excitability of rabbit CA1 pyramidal neurons in an age- and concentration-dependent manner. J Neurophysiol. 68(6):2100−9.
  212. Moyer J.R. Jr, Disterhoft J.F. (1994) Nimodipine decreases calcium action potentials in rabbit hippocampal CA1 neurons in an age-dependent and concentration-dependent manner. Hippocampus. 4(1):11−7.
  213. Moyer J.R. Jr, Thompson L.T., Disterhoft J.F. (1996) Trace eyeblink conditioning increases CA1 excitability in a transient and learning-specific manner.
  214. J Neurosci. 16(17): 5536−46.
  215. Moyer J.R. Jr, Power J.M., Thompson L.T., Disterhoft J.F. (2000) Increased excitability of aged rabbit CA1 neurons after trace eyeblink conditioning. J Neurosci. 20(14): 5476−82.
  216. R.M., Malenka R.C. (1992) Mechanisms underlying induction of homosynaptic long-term depression in area CA1 of the hippocampus. Neuron 9: 967−975.
  217. R.M., Herron C.E., Malenka R.C. (1993) An essential role for protein phosphatases in hippocampal long-term depression. Science. 261(5124): 1051−5.
  218. W.E., Gispen W.H. (1996) The current status of the calcium hypothesis of brain aging and Alzheimer’s disease. Life Sci. 59: 357−510.
  219. G.G., Fedorov N.B., Giese K.P. Ohno M., Friedman E., Chen R., Silva A.J. (2004) Increased neuronal excitability, synaptic plasticity, and learning in aged Kvbettal.l knockout mice. Curr. Biol. 14: 1907−1915.
  220. Nicoll R (1988) The coupling of neurotransmitter receptors to ion channels in the brain. Science. 241(4865): 545−51. Review.
  221. R., Malenka R.C., Kauer J.A. (1990) Functional comparison of neurotransmitter receptor subtypes in mammalian central nervous system. Physiol. Rev. 70(2): 513−565.
  222. C.M., Korol D.L., Foster T.C. (1996) Increased susceptibility to induction of long-term depression and long-term potentiation reversal during aging. J. Neurosci. 16: 5382−5392.
  223. C.M., Halpain S., Foster T.C. (1998) Reversal of age-related alterations in synaptic plasticity by blockade of L-type Ca2+ channels. J. Neurosci. 18: 3171−3179.
  224. C.M., Foster T.C. (1999) MK-801 improves retention in aged rats: implications for altered neural plasticity in age-related memory deficits. Neurobiol. Learn. Mem. 71: 194−206. .
  225. O’Dell T.J., Kandel E.R., Grant S.G. (1991) Long-term potentiation in the hippocampus is blocked by tyrosine kinase inhibitors. Nature 353: 558−560.
  226. Oh M.M., Power J.M., Thompson L.T., Moriearty P.L., Disterhoft J.F. (1999) Metrifonate increases neuronal excitability in CA1 pyramidal neurons from both young and aging rabbit hippocampus. J Neurosci. 19(5): 1814−23.
  227. Oh M.M., Gamelli A.E., Wu W.W., Sametsky E., Disterhoft J.F. (2001) Morris watermaze learning enhances neuronal excitability of CA1 hippocampal pyramidal neurons in rats. Soc. Neurosci. Abstr. 27: 921.1.
  228. Ohl F., Roedel A., Binder E., Holsboer F. (2003) Impact of high and low anxiety on cognitive performance in a modified hole board test in C57BL/6 and DBA/2 mice.
  229. Eur J Neurosci. 17(1): 128−36.
  230. Y., Ishibashi T., Okada K., Ishida K., Nakamura M. (2005) Trace eyeblink conditioning requires the hippocampus but not autophosphorilation of {alpha} CaMII in mice. Learn. Mem. 12:211−215.
  231. M., Tseng W., Silva A.J., Disterhoft J.F. (2005) Trace eyeblink conditioning requires the hippocampus but not autophosphorylation of {alpha}CaMKII in mice. Learn. Mem. 12:211−215.
  232. J.A., Markus E.J. (1998) Age-related deficits non the radial maze and in fear conditioning: hippocampal processing and consolidation. Hippocampus 8: 402−415.
  233. M.W., Kessler M., Larson J., Schottler F., Lynch G. (1990) Glycine site associated with the NMDA receptor modulates long-term potentiation. Synapse 5:265 270.
  234. N.A., Lisman J.E. (1998) D1/D5 dopamine receptors inhibit depotentiation at CA1 synapses via cAMP-dependent mechanism. J Neurosci. 18(4): 1270−9.
  235. A., Zhang L., Charlton M.P., Carlen P.L. (1999) Differential modulation of synaptic transmission by calcium chelators in young and aged hippocampal CA1 neurons: evidence for altered calcium homeostasis in aging. J Neurosci. 19(3): 906−15.
  236. A., Nogues X., Marighetto A., Micheau J., Battaini F., Govoni S., Jaffard R. (1998) Cytosolic hippocampal PKC and aging: correlation with discrimination performance. Neuroreport. 9(4): 725−9.
  237. P., Storm J.F. (1993) PKA mediates the effects of monoamine transmitters on the K+ current underlying the slow spike frequency adaptation in hippocampal neurons. Neuron. 11(6): 1023−35.
  238. P., Storm J.F. (1995) Dopamine modulates the slow Ca(2+)-activated K+ current IAHP via cyclic AMP-dependent protein kinase in hippocampal neurons.
  239. J Neurophysiol. 74(6): 2749−53.
  240. P., Storm J.F. (1996) Interaction between alpha- and beta-adrenergic receptor agonists modulating the slow Ca(2+)-activated K+ current IAHP in hippocampal neurons. Eur J Neurosci. 8(10): 2098−110.
  241. P., Krause M., Haug T., Storm J.F., Stuhmer W. (1998) Modulation of the Ca2±activated K+ current sIAHP by a phosphatase-kinase balance under basal conditions in rat CA1 pyramidal neurons. J. Neurophysiol. 79: 3252−3256.
  242. M.A. Smith M.Y. Gallagher M. (1987) Spatial learning impairments in aged rats trained with a salient configuration of stimuli. Psychobiology 15:248−254.
  243. T.A., Landfield P.W. (1990) Aging-related prolongation of calcium spike duration in rat hippocampal slice neurons. Brain Res. 508(1): 1−6.
  244. Porter NM, Thibault O, Thibault V, Chen KC, Landfield PW (1997) Calcium channel density and hippocampal cell death with age in long-term culture. J Neurosci 17:5629−5639.
  245. Power J.M., Oh M.M., Disterhoft J.F. (2001) Metrifonate decreases sI (AHP) in CA1 pyramidal neurons in vitro. J Neurophysiol. 85(1): 319−22.
  246. Power J.M., Wu W.W., Sametsky E., Oh M.M., Disterhoft J.F. (2002) Age-related enhancement of the slow outward calcium-activated current in hippocampal CA1 pyramidal neurons in vitro. J. Neuroscience 22(16) 7234−7243.
  247. R.J., Milgram S., Hafner S. (1983) Long-term potentiation phenomena in the rat limbic forebrain. Brain. Res. 260: 217−231.
  248. Rapp P. R, Rosenberg R.A. Gallagher M. (1987) An evaluation of spatial information processing in aged rats. Behav. Neurosci. 101: 3−12.
  249. P. R. (1993) Neuropsychological analysis of learning and memory in the aged nonhuman primate. Neurobiol. Aging 14(6): 627−629.
  250. P.H., Levitan I.B. (1995) Kinase and phosphatase activities intimately associated with a reconstituted calcium-dependent potassium channel.
  251. J Neurosci. 15(6): 4572−9.
  252. Reymann K.G., Malish R., Schulzeck K., Brodemann R., Ott T., Matties H. (1985) The duration of long-term potentiation in the CA1 region of the hippocampal slice preparation. Brain Res. Bull. 15: 249−255.
  253. Rosenzweig E.S., Rao G., McNaughton B.L., Barnes C.A. (1997) Role of temporal summation in age-related LTP induction deficits. Hippocampus 7: 549−558.
  254. E.S., Barnes C.A. (2003) Impact of aging on hippocampal function: plasticity, network dynamics, and cognition. Prog Neurobiol. 69(3): 143−79. Review.
  255. D., Grossman Y., Barkai E. (1998) Reduced after-hyperpolarization in rat piriform cortex pyramidal neurons is associated with increased learning capability during operant conditioning. Eur J Neurosci. 10(4): 1518−23.
  256. Sah P. (1996) Ca (2+)-activated K+ currents in neurones: types, physiological roles and modulation. Trends Neurosci. 19(4): 150−4. Review.
  257. Sah P., Isaacson J.S. (1995) Channels underlying the slow afterhyperpolarization in hippocampal pyramidal neurons: neurotransmitters modulate the open probability. Neuron. 15(2): 435−41.
  258. Sah P., Bekkers J.M. (1996) Apical dendritic location of slow afterhyperpolarization current in hippocampal pyramidal neurons: implications for the integration of long-term potentiation. J Neurosci. 16(15): 4537−42.
  259. T. (1977) Central neurotransmitter substances and aging: a review. J Am Geriatr Soc. (8): 337−48. Review.
  260. T.J. (1977) Storing covariance with nonlineary interacting neurons. J. Math. Biol. 4:303−321.
  261. S. Teyler T.J., Robbins N. (1998) Aging differently alters forms of long-term potentiation in rat hippocampal area CA1. J. Neurophysiol. 79: 334−341.
  262. Sharp P.E., Barnes C.A., McNaughton B.L. (1987) Effects of aging on environmental modulation of hippocampal evoked responses. Behav Neurosci. 101(2): 170−8.
  263. J., Barnes C.A. (1996) Age-related decrease in cholinergic synaptic transmission in three hippocampal subfields. Neurobiol. Aging 17:439−451.
  264. Shen J., Barnes C.A., McNaughton B.L., Skaggs W.E., Weaver K.L. (1997) The effect of aging on experience-dependent plasticity of hippocampal place cells J. Neurosci. 17: 6769−6782.
  265. Shi S.H., Hayashi Y., Petralia R.S., Zaman S.H., Wenthold R.J., Svoboda K., Malinow R. (19 999) Rapid spine delivery and redistribution of AMPA receptors after synaptic NMDA receptor activation. Science 284: 1811−1816.
  266. S., Kawasaki K. (1997) Electrophysiological changes in rat hippocampal pyramidal neurons produced by cholecystokinin octapeptide. Neuroscience. 78(4): 100 516.
  267. Schurman T, Traber J (1994) Calcium antagonists in aging brain. Ann N Y Acad Sci 747:467−474.
  268. V.G., Vorobyev V.S. (1979) A study of synaptic plasticity in hippocampal slices. Acta Neurobiol Exp (Wars). 39(6):633−42.
  269. V.G., Chepkova A.N. (1998) Hebbian synapses in cortical and hippocampal pathways. Rev Neurosci. 9(4):243−64. Review.
  270. Solomon PR, Wood MS, Groccia-Ellison ME, Yang BY, Fanelli RJ, Mervis RF (1995) Nimodipine facilitates retention of the classically conditioned nictitating membrane response in aged rabbits over long retention intervals. Neurobiol Aging 16:791−796.
  271. Sourdet V, Russier M, Daoudal G, Ankri N, Debanne D (2003) Long-term enhancement of neuronal excitability and temporal fidelity mediated by metabotropic glutamate receptor subtype 5. J Neurosci 23:10 238−10 248.
  272. L.R. (1987) Memory and brain, Oxford Univ. Press, New York. P. 25.
  273. O. (1976) Topographic organization of the projection from the enthorinal are to the hippocampal formation of the rat. J. Comp. Neurol., 167(3): 287−314.
  274. Teyler T.J., Cavus I., Coussens C., DiScenna P., Grover L., Lee Y.P., Little Z. (1994) Multideterminant role of calcium in hippocampal synaptic plasticity. Hippocampus. 4(6):623−34. Review.
  275. Tielen A.M., Mollevagner W.J., Lopes da Silva F.H., Hollander C.F. (1983) Neuronal plasticity in hippocampal slices of extremely old rats. In: Gispen, W.H., Trber J. (Eds.), Aging of the Brain. Elsevier, Amsterdam, pp. 73−84.
  276. O., Landfield P.W. (1996) Increase in single L-type calcium channels in hippocampal neurons during aging. Science. 272(5264): 1017−20.
  277. O., Porter N.M., Chen K.C., Blalock E.M., Kaminker P.G., Clodfelter G.V., Brewer L.D., Landfield P.W. (1998) Calcium dysregulation in neuronal aging and Alzheimer’s disease: history and new directions. Cell Calcium. 24(5−6): 417−33. Review.
  278. Thibault O, Hadley R, Landfield PW (2001) Elevated postsynaptic Ca2+.i and L-type calcium channel activity in aged hippocampal neurons: relationship to impaired synaptic plasticity. J Neurosci 21:9744−9756.
  279. E., Norman E.D., Barrionuevo G., Klann E. (1998) Transient and persistent increases in protein phosphatase activity during long-term depression in the adult hippocampus in vivo. Neuroscience. 86(4): 1023−9.
  280. K.L., Davis S., Hunt S.P., Laroche S. (1996) Alterations in the expression of specific glutamate receptor subunits following hippocampal LTP in vivo.1.arn Mem. 3(2−3): 197−208.
  281. Thompson L.T., Moyer J.R. Jr., Disterhoft J.F. (1996a) Transient changes in excitability of rabbit CA3 neurons with a time course appropriate to support memory consolidation. J. Neurophysiol. 76: 1836−1849.
  282. Thompson L.T., Moyer J.R. Jr, Disterhoft J.F. (1996b) Trace eyeblink conditioning in rabbits demonstrates heterogeneity of learning ability both between and within age groups. Neurobiol Aging. 17(4): 619−29.
  283. L.T., Disterhoft J.F. (1997) Age- and dose-dependent facilitation of associative eyeblink conditioning by D-cycloserine in rabbits. Behav. Neurosci. Ill: 1303−1312.
  284. G.E., Chaput Y., Andrade R. (1995) Cyclic AMP and protein kinase A mediate 5-hydroxytryptamine type 4 receptor regulation of calcium-activated potassium current in adult hippocampal neurons. Mol Pharmacol. 47(1): 191−7.
  285. G.E., Arfken C.L., Andrade R. (1996) 5-Hydroxytryptamine4 receptors reduce afterhyperpolarization in hippocampus by inhibiting calcium-induced calcium release. Mol Pharmacol. 50(5): 1316−22.
  286. A., Toescu E.C. (1998) Calcium and neuronal ageing. Trends Neurosci. 21(l):2−7. Review.
  287. J., Dunnett C., Falls W.A. (2004) C57BL/6J and DBA/2J differ in extinction and renewal of extinguished conditional fear. Behav. Brain Res. 154(2): 56 776.
  288. Y.T., Salter M.W. (1994) Regulation of NMDA receptors by tyrosine kinases and phosphatases. Nature. 369(6477): 233−5.
  289. Wang L.Y., Orser B.A., Brautigan D.L., MacDonald J.F. (1994) Regulation of NMDA receptors in cultured hippocampal neurons by protein phosphatases 1 and 2A. Nature. 369(6477):230−2.
  290. J.H., Kelly P.T. (1996) The balance between postsynaptic Ca(2+)-dependent protein kinase and phosphatase activities controlling synaptic strength.1.arn Mem. 3(2−3): 170−81.
  291. Y., Rowan M.J., Anwyl R. (1997) LTP induction dependent on activation of Ni2±sensitive voltage-gated calcium channels, but not NMDA receptors, in the rat dentate gyrus in vitro. J. Neurophysiology. 78(5): 2574−81.
  292. G. (1988) A neuropsychological analysis of memory loss with age. Neurobiol. Aging 9: 487−494.
  293. Wu WW, Chan CS, Disterhoft JF (2004) Slow afterhyperpolarization governs the development of NMDA receptor-dependent afterdepolarization in CA1 pyramidal neurons during synaptic stimulation. J Neurophysiol 92:2346−2356.
  294. D.J., Nicoll R.A. (1994) A role for protein kinases and phosphatases in the Ca(2+)-induced enhancement of hippocampal AMPA receptor-mediated synaptic responses. Neuron. 13(3):635−43.
  295. Xu J., Kang N., Jiang L., Nedergaard M., Kang J. (2005) Activity-dependent long-term potentiation of intrinsic excitability in hippocampal CA1 pyramidal neurons. J. Neurosci. 25:1750−1760.
  296. C. Mcllwain H. (1966) Electrical activities in thin sections from the mammalian brain maintained in chemically defined media in vitro. J. Neurochemistry 13: 1333−1343.
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