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/Subject (In\0402008\040we\040published\040the\040first\040set\040of\040guidelines\040for\040standardizing\040research\040in\040autophagy\056\040Since\040then\054\040research\040on\040this\040topic\040has\040continued\040to\040accelerate\054\040and\040many\040new\040scientists\040have\040entered\040the\040field\056\040Our\040knowledge\040base\040and\040relevant\040new\040technologies\040have\040also\040been\040expanding\056\040Accordingly\054\040it\040is\040important\040to\040update\040these\040guidelines\040for\040monitoring\040autophagy\040in\040different\040organisms\056\040Various\040reviews\040have\040described\040the\040range\040of\040assays\040that\040have\040been\040used\040for\040this\040purpose\056\040Nevertheless\054\040there\040continues\040to\040be\040confusion\040regarding\040acceptable\040methods\040to\040measure\040autophagy\054\040especially\040in\040multicellular\040eukaryotes\056\040A\040key\040point\040that\040needs\040to\040be\040emphasized\040is\040that\040there\040is\040a\040difference\040between\040measurements\040that\040monitor\040the\040numbers\040or\040volume\040of\040autophagic\040elements\040\050e\056g\056\054\040autophagosomes\040or\040autolysosomes\051\040at\040any\040stage\040of\040the\040autophagic\040process\040vs\056\040those\040that\040measure\040flux\040through\040the\040autophagy\040pathway\040\050i\056e\056\054\040the\040complete\040process\051\073\040thus\054\040a\040block\040in\040macroautophagy\040that\040results\040in\040autophagosome\040accumulation\040needs\040to\040be\040differentiated\040from\040stimuli\040that\040result\040in\040increased\040autophagic\040activity\054\040defined\040as\040increased\040autophagy\040induction\040coupled\040with\040increased\040delivery\040to\054\040and\040degradation\040within\054\040lysosomes\040\050in\040most\040higher\040eukaryotes\040and\040some\040protists\040such\040as\040Dictyostelium\051\040or\040the\040vacuole\040\050in\040plants\040and\040fungi\051\056\040In\040other\040words\054\040it\040is\040especially\040important\040that\040investigators\040new\040to\040the\040field\040understand\040that\040the\040appearance\040of\040more\040autophagosomes\040does\040not\040necessarily\040equate\040with\040more\040autophagy\056\040In\040fact\054\040in\040many\040cases\054\040autophagosomes\040accumulate\040because\040of\040a\040block\040in\040trafficking\040to\040lysosomes\040without\040a\040concomitant\040change\040in\040autophagosome\040biogenesis\054\040whereas\040an\040increase\040in\040autolysosomes\040may\040reflect\040a\040reduction\040in\040degradative\040activity\056\040Here\054\040we\040present\040a\040set\040of\040guidelines\040for\040the\040selection\040and\040interpretation\040of\040methods\040for\040use\040by\040investigators\040who\040aim\040to\040examine\040macroautophagy\040and\040related\040processes\054\040as\040well\040as\040for\040reviewers\040who\040need\040to\040provide\040realistic\040and\040reasonable\040critiques\040of\040papers\040that\040are\040focused\040on\040these\040processes\056\040These\040guidelines\040are\040not\040meant\040to\040be\040a\040formulaic\040set\040of\040rules\054\040because\040the\040appropriate\040assays\040depend\040in\040part\040on\040the\040question\040being\040asked\040and\040the\040system\040being\040used\056\040In\040addition\054\040we\040emphasize\040that\040no\040individual\040assay\040is\040guaranteed\040to\040be\040the\040most\040appropriate\040one\040in\040every\040situation\054\040and\040we\040strongly\040recommend\040the\040use\040of\040multiple\040assays\040to\040monitor\040autophagy\056\040In\040these\040guidelines\054\040we\040consider\040these\040various\040methods\040of\040assessing\040autophagy\040and\040what\040information\040can\054\040or\040cannot\054\040be\040obtained\040from\040them\056\040Finally\054\040by\040discussing\040the\040merits\040and\040limits\040of\040particular\040autophagy\040assays\054\040we\040hope\040to\040encourage\040technical\040innovation\040in\040the\040field\056)
/Keywords 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