ﻢﻫد ﺰﻴﺳ هرﺎﻤ ﺷ ﻢﺘﻔ ﻫ لﺎﺳ تﻼﻀﻋ ﻲﻓﺮﺼﻣ ﺰﻛﻮﻠﮔ و ﺎﻤﺳﻼﭘ دازآ بﺮ
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1 رابطه بين اسيد چرب آزاد پلاسما و گلوكز مصرفي عضلات مصطفي جانجان دانشجو تربيتبدني و علوم ورزشي دانشگاه غيرانتفاعي شمال آمل سيده ندا علوي دانشجو تربيتبدني و علوم ورزشي دانشگاه غيرانتفاعي شمال آمل چكيده با توجه به اهميت گلوكز به عنوان سوخت ويژه ي مغز استفاده از چربي به جاي كربوهيدرات در صورت امكان هنگام فعاليت ورزشي اهميت زيادي دارد.هنگام فعاليت ورزشي استقامتي فرآيند ليپوليز افزايش مي يابد. اين عمل عمدتا بر اثر افزايش اپي نفرين موجود در خون افزايش فعاليت اعصاب سمپاتيكي ميباشد. بالا رفتن اسيد چرب آزاد (FFA) پلاسما سبب افزايش غلظت استيل- كوآ و NADH در ماتريكس ميتوكندريايي مي شوند كه در ابتدا باعث مهار واكنش پيروات دهيدروژناز مي شود.هدف از انجام مطالعه مورد نظر بررسي رابطهي بين افزايش اسيد چرب آزاد پلاسما بر حمل و نقل گلوكز و ميزان مصرف آن در عضلات مي باشد. به همين منظور اطلاعاتي مربوط به اثرات اسيد چرب بر انتقال گلوكز از مطالعات انجام شده قبلي جمعآوري شد. مقالات كار شده در اين زمينه را به دو گروه موافق و مخالف دستهبندي و مورد بررسي قرار داديم. نتايج حاصل از بررسي مطالعات نشان داد كه افزايش اسيد چرب آزاد پلاسما سبب كاهش گلوكز مصرفي عضلات مي شود و همچنين اين افزايش سبب كاهش انتقال گلوكز به عضلات مي شود. واژگان كليدي: ورزش استقامتي ليپوليز سمپاتيكي NADH FFA 53
2 مقدمه متابوليسم كه شامل همه واكنشهاي شيميايي يا فرايند هايي است كه در ارگانيزم انجام ميشود تا حد زيادي مثل موتور بنزيني عمل مي كند. موتور براي آنكه كار مفيدي انجام دهد به اكسايش مواد سوختي وابسته است. در سلول هايي كه واكنشهاي استفاده از ATP به كندي انجام مي شود غلظت ATP را مي توان به آساني و با افزايش تدريجي واكنشهايي كه ATP توليد مي كنند مثل اكسيداسيون مواد سوختي ثابت نگه داشت. اين مواد سوختي شامل كربوهيدراتها چربي ها پروتي ين ها و... هستند.گلوكز از مهمترين مواد سوختي است. كه مي تواند انرژي زيادي را در اختيار ما قرار دهد.در فعاليت هاي بي هوازي سوخت غالب گلوكز مي باشد و سبب توليد اسيد لاكتيك و انرژي مي شود. از ديگر مواد سوختي چربيها هستند. بيشتر افراد به خوبي مي دانند كه چربيها منابع سوختي مهمي در بدن اند و در سرتاسر بدن ذخيره مي شوند. چربي به صورت تري آسيل گليسرول در سلولهاي چربي ذخيره مي شود. چربي ذخيره انرژي دراز مدت است در حالي كه گليكوژن ذخيره ي انرژي كوتاه مدت به شمار مي رود. از آنجايي كه اسيدهاي چرب در آب نامحلولاند در خون به پروتي ين آلبومين متصل و تحت عنوان اسيدهاي چرب آزاد (FFA) در خون حمل مي شوند. در فعاليت هوازي به علت وجود اكسيژن كافي در بدن ميزان كاتابوليسم اسيدهاي چرب افزايش پيدا مي كند. با توجه به اهميت گلوكز به عنوان سوخت ويژه ي مغز استفاده از چربي و كربوهيدرات در صورت امكان هنگام فعاليت ورزشي اهميت زيادي دارد. هنگام فعاليت ورزشي استقامتي فرآيند ليپوليز افزايش مي يابد. اين عمل عمدتا بر اثر افزايش اپي نفرين موجود در خون افزايش فعاليت اعصاب سمپاتيكي است. در شروع فعاليت ورزشي زير بيشينه ميزان برداشت FFA توسط عضلات فعال افزايش مييابد. اين FFA برداشتي بسيار بيشتر از آن FFA است كه در اين لحظات از ليپوليز بافت چربي بدست مي آيد به طوري كه اسيدهاي چرب پلاسما در ابتدا كاهش مي يابند. با افزايش تدريجي ليپوليز ميزان رهايش FFA از بافت چربي از ميزان FFA برداشتي از پلاسما پيشي مي گيرد. در نتيجه غلظت FFAپلاسما به افزايش بيشتر در اكثر دوره زماني فعاليت ورزشي تمايل دارد. بالا رفتن FFA پلاسما سبب افزايش غلظت استيل كوآ و NADH در ماتريكس ميتوكندريايي ميشوند كه در ابتدا باعث مهار واكنش پيروات دهيدروژناز مي شود.هنگامي كه غلظت ATP 54
3 NADH يا استيل كوآنزيم A بالا باشد. پيروات دهيدروژناز به وسيله آنزيمي به نام پيروات دهيدروژناز كيناز فسفريله مي شود. در اثر فسفوريلاسيون فعاليت آنزيم كاهش مي يابد. افزايش غلظت استيل كوآ در ماتريكس به افزايش سيترات در ماتريكس منجر مي شود. بنابراين سيترات سيتوزولي افزايش مي يابد. فسفوفروكتوكيناز (PFK) يكي از آنزيم هاي محدود كننده اين مسير ميباشد. اين آنزيم به طور آلوستريك بوسيله ATP مهار و توسط AMP تحريك مي شود. فسفوفروكتوكيناز باعث افزايش فروكتوز 1 و 6 دي فسفات مي شود. فروكتوز 1 و 6 دي فسفات در نهايت به فسفوانول پيروات تبديل و باعث افزايش استيل كوآنزيم و سيترات و بالا رفتن ميزان ATP مي شود. همانطور كه بيان شد ATP سبب مهار PFK مي شود. بنابراين سيترات سبب مهار فسفو فروكتوكيناز شده و ايزوآنزيمهاي هگزوكيناز III,II,I از طريق محصول واكنش يعني گلوكز 6 -فسفات مهار ميشوند و در نتيجه گلوكز فسفوريله نخواهد شد و غلظت آن در سلول افزايش مي يابد.اين افزايش شيب انتقال گلوكز به داخل سلول را كاهش مي دهد و در نتيجه ورود گلوكز به داخل سلول را كند مي كند. براي متاليزه شدن گلوكز در درون سلول دو فرآيند بايد رخ دهد: گلوكز بايد با حامل گلوكز به درون سلول انتقال يابد و سپس به گلوكز 6 -فسفات فسفوريله شود.اينكه كداميك از اين دو فرايند مرحله بعدي متابوليسم گلوكز را محدود مي كند مورد بحث و جدل است. با وجود اين تحت شرايط فيزولوژيك طبيعي تر غلظت گلوكز آزاد موجود در درون سلولهاي عضله اسكلتي بسيار كم است.اين امر به اين نكته اشاره دارد كه انتقال گلوكز عامل محدود كننده اي براي متابوليسم گلوكز است زيرا گلوكز تقريبا با همان سرعتي كه فسفوريله مي شود وارد سلول مي گردد.افزايش چشمگير FFA خون مصرف كلي كربوهيدرات و نيز كربوهيدرات مصرفي عضله اسكلتي را در افراد در حال استراحت كاهش ميدهد. براساس مطالعههاي جديدتر به نظر ميرسد افزايش يكبارهي FFA ناشي از تزريق ليپيدها به درون جريان خون اكسيداسيون كربوهيدرات را در چند نقطه كاهش مي دهد. اگر غلظت FFA تا چند ساعت زياد باشد احتمالا برداشت گلوكز از طريق كاهش پاسخ انتقال دهنده هاي گلوكز (GLUT-4) به گلوكز تضعيف مي- شود كه دليل آن وجود FFA زياد است. همچنين افزايش FFA غلظت فعال كنندههاي آلوستريك در سيتوزول تار عضلاني (pi,amp) را كاهش مي دهد. اگر جريان از مسير گليكوليتيكي به پيروات يا از طريق كاهش برداشت گلوكز يا مهار فسفوفروكتوكيناز تضعيف شود دسترسي به پيروات براي واكنش پيروات و هيدروژناز در ماتريكس كمتر مي شود.در مجموع كربوهيدرات كاهش 55
4 مي يابد كه افزايش دسترس به استيل كوآي ناشي از اكسيد ليپيدها جبران مي شود.هدف از اين مطالعه بررسي اثر FFA پلاسما بر انتقال گلوكز و ميزان مصرف آن در عضلات مي باشد. بحث و نتيجهگيري كاستيل و همكاران (1997) اثر افزايش FFA پلاسما در استفاده از گليكوژن عضله را در هفت آزمودني مورد بررسي قرار دارند. هفت مورد به مدت 30 دقيقه روي تردميل با روش تقريبي %70 vo 2 max فعاليت كردند. اين افزايش FFA پلاسما موجب كاهش %40 ميزان تخليه گليكوژن عضله شد.اين داده ها نشان مي دهد كه در طول فعاليت شديد و طولاني عضلاني انسولين پلاسما و FFA هر دو در ميزان اكسيداسيون كربوهيدرات ها تاثير دارند. رينه و همكاران (1976) اظهار كردند كه افزايش اكسيداسيون اسيدهاي چرب آزاد باعث مهار مصرف گليكوژن توسط عضله مي شود كه احتمالا اين فراخواني FFA سبب مهار ميزان بهره برداري عضله از گليكوژن بوسيله عمل انسولين و قند خون و در طولاني مدت سبب كاهش شتاب فعاليت عضلاني مي شود. درسنر و همكاران دادند نشان (1998) كه افزايش غلظت فسفوريلاسيون ميشود و دلالت بر مهار FFA (1994) در مورد افزايش FFA عمل انتقال باعث كاهش انتقال قند خون يا فعاليت دارد. در تحقيق ديگري كه بودن و همكاران پلاسما و كاهش تحريك انسولين و جذب گلوكز انجام راند به نتايج قابل توجهي دست يافتند. آنها نتيجه گرفتند كه افزايش FFA باعث مهار جذب گلوكز و تحريك انسولين توسط سنتز گليكوژن و كاهش اكسيداسيون كربوهيدرات مي شود و اين افزايش FFA و يا گليسرول سبب سركوب انسولين كبدي و خروجي گلوكز و در نتيجه باعث مقاومت به انسولين مي شود.در اين تحقيق دو نقص را متوجه مي شويم كه عبارتند از: نقص اختلاف در فعاليت گليكوژن فسفاتاز. اين اختلال با افزايش گلوكز 6 فسفات همراه مي شود. نقص دوم كه قبل از نقص گليكوژن فسفاژ مشاهده مي شود افزايش غلظت FFA باعث كاهش گلوكز 6 فسفات عضله مي شود.افزايش FFA و كاهش گلوكز 6 فسفات عضله مي تواند در اثر كاهش حمل و نقل گلوكز يا فعاليت فسفوريلاسيون باشد. پيي ل و همكارانش بيان كردند كه تمرين استقامتي درازمدت باعث افزايش غلظت گليكوژن مي شود. آنها غلظت گليكوژن عضله پاهاي تمرين كرده و تمرين نكرده چهار دوچرخه سوار را سنجيدند. مقادير گليكوژن عضله در پاي تمرين كرده بيشتر از پاي تمرين نكرده بوده است. هيكنر و همكاران (1997) ميزان سنتز گليكوژن را در 56
5 آزمودنيهاي تمرين كرده و تمرين نكرده پس از 6 و 72 ساعت بازيافت مقايسه كرده اند. فعاليت پيش از بازيافت هر دو گروه 2 ساعت با %75 max vo 2 بوده كه با پنج دوي سرعت يك دقيقه اي به اتمام رسيد. ميزان سنتز گليكوژن در آزمون ها تمرين كرده پس از 6 ساعت بازيافت بيشتر از دو برابر افراد تمرين نكرده است.اين پاسخ با افزايش سه برابري مقدار GLUTS عضله همراه است. رابرگز و اسكات اشاره كردهاند كه تمرينهاي استقامتي درازموت فعاليت دو آنزيم اصلي تنظيم كننده مسيرهاي گليكوژنوليز و گليكوليز يعني فسفوفروكتوكيناز (PFK) را افزايش مي دهند.اين نتيجه گيري كه نتايجي برخلاف موارد مورد بررسي را نشان مي دهد احتمالا به علت نوع پروتكل سنجيده شده دارد.در تمامي مقالات اندازه گيري ها در هنگام فعاليت سنجيده شده ولي در كتاب اثر فعاليت دراز مدت سنجيده است. راندل و همكارانش چرخه گلوكز- اسيد چرب را پيشنهاد كردند كه براساس آن اسيدهاي چرب اكسيداسيون كربوهيدراتها را در آماده سازي عضله قلب و ديافراگم كاهش مي دهند.تحقيقات نشان مي دهند كه فعاليت هاي استقامتي سبب بالا رفتن ميزان ليپوليز بافت چربي مي شود. اين افزايش ليپوليز سبب افزايش ميزان رهايش FFA از بافت چربي مي شود.بالا رفتن FFA پلاسما سبب مهار پيروات دهيدروژناز و در نهايت سبب كاهش گلوكز مصرفي عضلات و حمل ونقل آنها به عضلات مي شود. به همين جهت بر فعاليت بي هوازي بعد از آن اثر منفي دارد كه اين اثر بسته به فاصله ي بين فعاليت دارد. 57
6 منابع 1) Costill D. L., E. Coyle, G. Dalsey, W. Evaps, W. Fink, and D. Hoopes. (1977). Effects of elevated plasma FFA and insulin on muscle glycogen usage during exercise. 2) Rennie, M., Winder M., and Holloszy J. O. (1976). Asparagus effect of increased free fatty acids on muscle glycogen content in exercising rats. 3) Dresner A., et al., (1998). Effects of free fatty acids on glucose transport and IRS-1-assocated phosphatide inositol 3 kinase activity. 4) Boden G., et al., (1994). Mechanisms of fatty acid induced inhibition of glucose uptake. 5) Piehl k., Adolfsson s., and nazar k. (1974). Glycogen storage and glycogen syntheses activity in trained and untrained muscle of man. 6) Hikener R. C. et al., (1997). Muscle glycogen accumulation after endurance exercise in trained and untrained individuals. 7) Ferrannini E., Eugene J. Barrett, Stefano Bevilacqua, and Ralph A. (1983). Effect of fatty acids on glucose production and utilization in man. 8) Brons Charlotte, et al., (2009). Impact of short-term high-fat feeding on glucose and insulin metabolism in young healthy men. 9) Randle P. J., et al., (1963). The glucose fatty- acids cycle: its role in insulin sensitivity and the metabolic disturbances of diabetes mellitus. 9) گاي يني عباسعلي. (1386). بيوشيمي فغاليت بدني. انتشارات پيام نور. 10) پاسالار. (1385). چكيده بيوشيمي. انتشارات دانشگاه تهران. 11) شادان فرخ. (1384). فيزيولوژي پزشكي. انتشارات چهر. 12) گاي يني عباسعلي و همكاران. (1384). اصول بنيادي فيزيولوژي ورزشي. 58
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