Antioxidant potential of lichen species and their secondary metabolites. A systematic review

Figures & data

Figure 1. Chemical structures of depside atranorin (a), dibezofuran usnic acid (b), depsidone protocetraric acid (c), and pulvinic acid (d).

Table 1. Antioxidant activities of lichen extracts.

Lichen species	Solvent	LP inhibition (IC50 or %INH)	DPPH (IC50 or %INH)	SO^.- RSA (IC50 or %INH)	Reducing power (A700 or IC50)	Phenolic/flavonoid content	Isolated compounds/ composition	Reference
Arthoniaceae Reichenb. ex Reichenb.
Arthothelium awastii Patw. & Makhija	ME	2.1–68.4% 15.7 μg/ml	2.6–68.4% 13.2 μg/ml	Not studied (NS)	NS	Different content depending on lichen culture conditions	NS	Verma et al. (2008b)
Catillariaceae Hafellner.
Toninia candida (Weber) Th.Fr.	ChE, ME, PE	ChE: 41.7 μg/ml ME: 46.5 μg/ml PE: 21.5 μg/ml	ChE :48.9 μg/ml ME: 51.5 μg/ml PE: 50.1 μg/ml	NS	ChE: 56. 7 μg/ml ME: 78.5 μg/ml PE: 51.5 μg/ml	ChE: 45.2 mg GA/g ME: 76.3 mg GA/g PE: 42.9 mg GA/g	Nortistic acid, stictic acid, usnic acid, protocetraric acid, atranorin	Manojlovic et al. (2012b)
	AcE	NS	115.8 μg/ml	221.5 μg/ml	0.055 (500 μg/ml)	49.8 μg PE/mg	Norstictic acid, stictic acid, protocetraricacid, usnic acid, atranorin	Ranković et al. (2012)
Cladoniaceae Zenker.
Cladonia amaurocraea (Flörke) Schaerer	AcE, DmE, EE, ME	AcE: 80.3% DmE, EE, ME: numeric value not reported (NVNR)	NVNR	NS	NS	AcE: 0.018 mg/g DmE: 0.016 mg/g EE: 0.005 mg/g ME: 0.174 mg/g	Atranorin, usnic acid	Singh et al. (2011)
Cladonia clathrata Ahti & L. Xavier	EE 70%	NS	50.2% 69.3 μg/ml	NS	NS	371.8 mg GA/g	NS	Silva et al. (2010)
Cladonia digitata (L.) Hoffm.	ME	9.8 %	NS	NS	0.085	13.7 mg GA/g	NS	Ranković et al. (2010b)
Cladonia fimbriata (L.) Fr.	ME	20.4%	NS	NS	0.114	26.4 mg GA/g	NS	Ranković et al. (2010b)
Cladonia foliacea (Huds.) Willd.	ME	NS	>1000 μg/ml	NS	NS	78.1 mg GA/g (28.2 mg Rutin/g)	NS	Mitrovic et al. (2011)
Cladonia furcata (Hudson) Schrad.	AcE, ME	NVNR	AcE: 471.3 μg/ml ME: >1000 μg/ml	NVNR ME > AcE	NVNR AcE > ME	NS	NS	Luo et al. (2009)
	AcE, ME, WE	NS	NVNR ME > AcE > WE	NVNR ME > AcE > WE	NVNR ME > AcE > WE	AcE: 12.1 μg PE ME: 52.7 μg PE WE: 5.8 μg PE	NS	Kosanic et al. (2011)
	AcE	NS	44.8%	23.9%	0.051	12.9 μg PE/mg (10.6 μg RE/mg)	NS	Ranković et al. (2011)
Cladonia mediterranea P.A.Duvign. & Abbayes	AcE, DmE, EE, ME	AcE: 73.7% DmE, EE, ME: NVNR	NVNR	NS	NS	AcE: 0.006 mg/g DmE: 0.001 mg/g EE: 0.008 mg/g ME: 0.009 mg/g	Usnic acid	Singh et al. (2011)
Cladonia rangiformis Hoffm	ChE, ME, WE	ChE: 48.3% ME: 1.6% WE: −11.5%	NS	NS	ChE: 0.083 ME: 0.115 WE: 0.034	ChE: 0.048 μg GA/ml ME: 0.010 μg GA/ml WE: 0.004 μg GA/ml	NS	Yücel et al. (2007)
Graphidaceae Dumort.
Graphis guimarana Vain.	ME 10%	NS	NS	19.7–23.2 μg/ml	NS	NS	NS	Behera et al. (2006a)
Graphis nakanishiana Patw. & C.R. Kulk.	ME 10%	NS	NS	11.2–14.2 μg/ml	NS	NS	NS	Behera et al. (2006a)
Graphis schizograpta Müll. Arg.	ME 10%	NS	NS	12.3–25.6 μg/ml	NS	NS	NS	Behera et al. (2006a)
Icmadophilaceae Triebel
Thamnolia vermicularis (Sw.) Schaer.	ME	2 mg/ml: 67.0%	2 mg/ml: 72.0% 0.2 mg/ml: 36.0%	NVNR	NVNR	NS	NS	Luo et al. (2006)
Lecanoraceae Körb.
Lecanora atra (Hudson) Ach.	AcE, ME, WE	NS	AcE: 94.7% ME: 93.3% WE: 93.2%	AcE: 84.5% ME, WE: NVNR	NVNR	AcE: 73.0 μg PE ME: 71.0 μg PE WE: 69.8 μg PE (AcE: 54.8 μg RE ME: 53.7 μg RE WE: 52.6 μg RE)	NS	Kosanić and Ranković (2011)
	AcE	NS	94.7%	84.5%	0.109	73.0 μg PE/mg (54.8 μg RE/mg)	NS	Ranković et al. (2011)
Lecanora muralis (Schreber) Rabenh.	AcE	NS	52.3%	33.6%	0.061	43.2 μg PE/mg (34.6 μg RE/mg)	NS	Ranković et al. (2011)
Lobariaceae Chevall.
Lobaria pulmonaria (L.) Hoffm.	ME, WE	ME: 87.5% WE: −17.1%	NS	NS	ME: 0.417 WE: 0.233	ME: 87.9 mg GA/g WE: 39.2 mg GA/g	NS	Odabasoglu et al. (2004)
Nephromataceae Wetm. ex J. C. David & D. Hawksw.
Nephroma parile (Ach.) Ach.	ME	3.6%	NS	NS	0.098	6.2 mg GA/g	NS	Ranković et al. (2010a)
Ochrolechiaceae R. C. Harris ex Lumbsch & I. Schmitt
Ochrolechia parella (L.) A.Massal.	ME	8.5%	NS	NS	0.077	5.6 mg GA/g	NS	Ranković et al. (2010b)
Ochrolechia tartarea (L.) A. Massal.	ME	26.6%	NS	NS	0.202	29.4 mg GA/g	NS	Ranković et al. (2010a)
Parmeliaceae Zenker
Bryoria fusfescens (Gyeln.) Brodo & D. Hawksw	ME, WE	ME: 12.6% WE: 18.4%	NS	NS	ME: 0.215 WE: 0.201	ME: 48.6 mg GA/g WE: 30.3 mg GA/g	NS	Odabasoglu et al. (2005)
Cetraria aculeata (Schreber) Fr.	AcE, ME	NVNR	AcE: >1000 μg/ml ME: >1000 μg/ml	NVNR AcE > ME	NVNR AcE > ME	NS	NS	Luo et al. (2009)
Cetraria fastigiata (Nyl.) Kärnefelt	AcE, DmE, EE, ME	AcE: 80.3% DmE, EE, ME: NVNR	NVNR	NS	NS	AcE: 0.019 mg/g DmE: 0.048 mg/g EE: 0.009 mg/g ME: 0.050 mg/g	Lichensterinic acid	Singh et al. (2011)
Cetraria islandica (L.) Ach.	WE	50 μg: 96.0% 100 μg: 99.0% 200 μg: 100.0% 500 μg: 100.0%	NVNR	NVNR	NVNR	0.039 μg PE/mg	NS	Gülçin et al. (2002)
	AcE, ME, WE	NS	NVNR	NVNR	NVNR	AcE: 25.0 μg PE ME: 38.0 μg PE WE: 18.2 μg PE (AcE: 7.7 μg RE, ME: 25.8 μg RE, WE: 1.4 μg RE)	NS	Kosanić and Ranković (2011)
Cetraria pinastri (Scop.) Gray	ME	48.8%	NS	NS	0.188	32.9 mg GA/g	NS	Ranković et al. (2010b)
Evernia prunastri (L.) Ach.	BE, DE, EE, ME	Crude wax: weak activity DE: activity EE: activity ME: activity	NS	NS	NS	NS	NS	Racine et al. (1980)
	ME	NS	727.7 μg/ml	NS	35.5 μg AA/g 0.6 μg T/g	18.2 μg GA/mg	NS	Stojanovic et al. (2010)
	ME	NS	>1000 μg/ml	NS	NS	80.7 mg GA/g (27.46 mg rutin/g)	NS	Mitrović et al. (2011)
	AcE	NS	663.1 μg/ml	1033.6 μg/ml	500 μg/ml: 0.029	34.1 μg PE/mg	Evernic acid, physodic acid, usnic acid, atranorin, chloroatranorin	Kosanić et al. (2013)
Flavocetraria nivalis (L.) Kärnefelt & A. Thell	AcE, DmE, EE, ME	NVNR	NVNR	NS	NS	AcE: 0.013 mg/g DmE: 0.012 mg/g EE: 0.007 mg/g ME: 0.162 mg/g	Usnic acid	Singh et al. (2011)
Flavoparmelia caperata (L.) Hale	ME	NS	347.2 μg/ml	NS	21.6 μg AA/g 19.3 μg T/g	11.9 μg GA/mg	NS	Stojanović et al. (2010)
	ME	NS	549.0 μg/ml	NS	NS	90.8 mg GA/g (33.6 mg rutin/g)	NS	Mitrović et al. (2011)
Hypogymnia physodes (L.) Nyl.	ME	NS	79.7 μg/ml	NS	96.9 μg AA/g 25.3 μg T/g	17.5 μg GA/mg	NS	Stojanović et al. (2010)
	ME	NS	45.6 μg/ml	NS	NS	141.6 mg GA/g (20.1 mg rutin/g)	NS	Mitrović et al. (2011)
	AcE, ME, WE	NS	AcE: 60.2% ME: 73.2% WE: 30.9%	NVNR ME > AcE > WE	NVNR ME > AcE > WE	AcE: 30.1 μg PE ME: 86.8 μg PE WE: 6.3 μg PE	NS	Kosanić et al. (2011)
Lethariella cashmeriana Krog	Hot WE	NS	NS	NS	Activity: 35.4	NS	Chlororubrocashmeriquinone Rubrocashmeriquinone	Kinoshita et al. (2010)
Lethariella sernanderi (Most.) Obermayer	Hot WE	NS	NS	NS	Activity: 163.9	NS	Canarione Rubrocashmeriquinone Chlororubrocashmeriquinone	Kinoshita et al. (2010)
Lethariella sinensis J.C.Wei & Y.M.Jiang	Hot WE	NS	NS	NS	Activity: 23.8	NS	7-Chlorocanarione Rubrocashmeriquinone Chlororubrocashmeriquinone	Kinoshita et al. (2010)
Parmelia caperata (L.) Ach.	AcE, ME, WE	NS	NVNR AcE > ME > WE	NVNR AcE > ME > WE	NVNR AcE > ME > WE	AcE: 40.4 μg PE ME: 15.3 μg PE WE: 14.9 μg PE	NS	Kosanić et al. (2011)
	AcE	NS	46.4%	61.5%	0.057	40.4 μg PE/mg (15.3 μg RE/mg)	NS	Kosanić et al. (2012a)
Parmelia centrifuga (L.) Ach.	ME	54.2%	NS	NS	0.375	49.8 mg GA/g	NS	Ranković et al. (2010a)
Parmelia crinita Ach.	ME	16.0%	NS	NS	0.172	12.8 mg GA/g	NS	Ranković et al. (2010b)
Parmelia pertusa Schaerer	AcE, ME, WE	NS	NVNR	NVNR Weak activity	NVNR Weak activity	AcE: 18.2 μg PE ME: 34.6 μg PE WE: 12.0 μg PE (AcE: 6.6 μg RE, ME: 18.2 μg RE, WE: 5.3 μg RE)	NS	Kosanić and Rankovic (2011)
Parmelia saxatilis (L.) Ach.	ME	25.0%	No activity	NS	NS	1.0% (w/w)	NS	Gülluce et al. (2006)
	ME, WE	Ferric thiocyanate method WE: 86.1% ME: 94.8% TBA test WE > ME	WE: 19.8% ME: 63.6%	WE: 83.0% ME: 80.0%	ME > WE	WE: 24.3 mg PE/mg ME: 55.1 mg PE/mg	Atranorin, chloroatranorin, salazinic acid	Özen and Kinalioglu (2008)
	AcE	NS	55.3%	35.3%	0.066	40.6 μg PE/mg (20.6 μg RE/mg)	NS	Kosanić et al. (2012a)
Parmelia sulcata Taylor	ME	NS	493.6 μg/ml	NS	25.3 μg AA/g 41.9 μg T/g	7.9 μg GA/g	NS	Stojanović et al. (2010)
	ME	NS	584.2 μg/ml	NS	NS	88.3 mg GA/g (44.9 mg rutin/g)	NS	Mitrović et al. (2011)
	AcE, ME, WE	NS	NVNR AcE > ME > WE	NVNR AcE > ME > WE	NVNR AcE > ME > WE	AcE: 38.2 μg PE ME: 25.1 μg PE WE: 9.6 μg PE	NS	Kosanić et al. (2011)
	AcE	NS	38.8%	33.8%	0.034	18.2 μg PE/mg (6.6 μg RE/mg)	NS	Kosanić et al. (2012a)
Parmotrema pseudotinctorum (Abbayes) Hale	ME	NS	500 μg/ml: 90.7% + honey: 81.5%	NS	NS	NS	Atranorin, lecanoric acid	Prashith Kekuda et al. (2009)
	ME	NS	62.1–89.4%	NS	0.265–0.776	NS	Atranorin, lecanoric acid	Praveen Kumar et al. (2010)
Parmotrema stuppeum (Taylor) Hale	BE, AcE	BE: 30.0–65.0% AcE: 35.0–68.0%	NS	NS	NS	NS	Methyl orsellinate, orselinic acid, atranorin, lecanoric acid	Jayaprakasha and Rao (2000)
Parmotrema tinctorum (Delise ex Nyl.) Hale	ME	3.6–71.3% 11.5 μg/ml	1.1–57.2% 13.6 μg/ml	NS	NS	Depends on lichen culture conditions	NDR	Verma et al. (2008b)
Platismatia glauca (L.) Culb. & C. Culb.	ME	27.0%	No activity	NS	NS	1.1 % (w/w)	NS	Güllüce et al. (2006)
Pseudoevernia furfuracea (L.) Zopf	ME, WE	ME: 26.4% WE: 48.2%	NS	NS	ME: 0.229 WE: 0.229	ME: 75.1 mg GA/g WE: 61.7 mg GA/g	NS	Odabasoglu et al. (2005)
	AcE, ME, WE	NS	AcE: 87.3% ME: 57.9% WE: 33.9%	NVNR	NVNR	AcE: 76.4 μg PE ME: 37.0 μg PE WE: 18.7 μg PE (AcE: 37.6 μg RE, ME: 21.0 μg RE, WE: 1.8 μg RE)	NS	Kosanić et al. (2011)
	BuE, DcmE, EaE, ME	BuE: No activity DcmE: 17.8% EaE: 23.8% ME: 17.2%	TLC spray	NS	NS	NS	Atraric acid, metil hematommate, methyl chlorohematommate	Güvenç et al. (2012)
Pseudoevernia furfuracea (L.) Zopf	AcE	NS	401.7 μg/ml	632.0 μg/ml	500 μg/ml: 0.058	76.4 μg PE/mg	Physodic acid, physodalic acid, atranorin, chloroatranorin, 3-hydroxiphysodic	Kosanić et al. (2013)
Pseudophebe pubescens (L.) M. Choisy	AcE, DmE, EE, ME	AcE: 82.4% DmE, EE, ME: NVNR	AcE: 51.8% DmE, EE, ME: NVNR	NS	NS	AcE: 0.017 mg/g DmE: 0.012 mg/g EE: 0.001 mg/g ME: 0.027 mg/g	Atranorin	Singh et al. (2011)
	AcE, ME	NVNR	AcE:>1000 μg/ml ME: >1000 μg/ml	NVNR ME > AcE	NVNR AcE > ME	NS	NS	Luo et al. (2009)
Usnea antarctica Du Rietz	AcE, ME	NVNR	AcE: 445.7 μg/ml ME: >1000 μg/ml	NVNR AcE = ME	NVNR AcE > ME	NS	NS	Luo et al. (2009)
Usnea aurantiacoatra (Jacq.) Bory	AcE, ME	NVNR	AcE: 642.6 μg/ml ME: 791.3 μg/ml	NVNR AcE = ME	NVNR AcE > ME	NS	NS	Luo et al. (2009)
Usnea barbata Motyka	AcE	NS	667.9 μg/ml	979.3 μg/ml	500 μg/ml: 0.019	31.3 μg PE/mg	Norstictic acid, usnic acid, atranorin, chloroatranorin	Ranković et al. (2012)
Usnea complanata (Müller Arg.) Motyka	WE, EE, EaE, ME	WE: 132.4 μg/ml EE: 125.0 μg/ml EaE: 157.9 μg/ml ME: 74.6 μg/ml	WE: 25.0 μg/ml EaE: 25.0 μg/ml EE: 22.9 μg/ml ME: 80.0 μg/ml	NS	NS	TLC technique	Usnic acid, psoromic acid	Behera et al. (2012)
Usnea florida (L.) Weber ex F.H.Wigg.	AcE, ME	ME: 15.0% WE: 13.4%	NS	NS	ME: 0.069 WE: 0.083	ME: 10.5 mg GA/g WE: 10.4 mg GA/g	NS	Odabasoglu et al. (2004)
Usnea ghattensis G. Awasthi	AcE, DmE, ME, PE	AcE: 55.0% DmE: 17.0% ME: 87.0% PE: 38.0%	AcE: 31.0% DmE: 11.0% ME: 73.0% PE: 43.0%	AcE: 18.0% DmE: 7.0% ME: 56.0% PE: 27.0%	NS	AcE: 14.0 mg/g DmE: 12.0 mg/g ME: 35.0 mg/g PE: 9.0 mg/g	NS	Behera et al. (2005a)
	ME	2–20 mg/ml: 3.8–73.3%	NVNR	20 mg/ml: 30.0%	NVNR	13.0 μg PE/mg	NS	Behera et al. (2006c)
	ME	67.0%	89.6%	89%	NS	NS	Usnic acid, nortistic acid	Verma et al. (2008a)
Usnea longissima Ach.	ME, WE	ME: 82.4% WE: 25.6%	NS	NS	ME: 0.178 WE: 0.100	ME: 38.6 mg GA/g WE: 18.3 mg GA/g	NS	Odabasoglu et al. (2004)
	ME	6 mg/ml: 97.3%	6 mg/ml: 1.0 mg/ml	NS	NS	2.62%	NS	Kim and Cho (2007)
Peltigeraceae Dumort.
Peltigera rufescens (Weiss) Humb.	ME, WE	ME: 74.0% WE: 26.4%	NS	NS	ME: 0.542 WE: 0.218	ME: 66.4 mg GA/g WE: 38.1 mg GA/g	NS	Odabasoglu et al. (2005)
Physciaceae Zahlbr.
Anaptychia ciliaris (L.) Körb.	ME	22.6%	NS	NS	0.195	27.6 mg GA/g	NS	Ranković et al. (2010a)
Heterodermia podocarpa (Bel.) D. D. Awasthi	ME	3.4–52.7% 12.7 μg/ml	1.6–53.7% 18.4 μg/ml	NS	NS	Different contents depending on lichen culture conditions	NDR	Verma et al. (2008b)
Physcia caesia (Hoffm.) Fürnr.	AcE, DmE, EE, ME	NVNR	NVNR	NS	NS	AcE: 0.065 mg/g EE: 0.015 mg/g ME: 0.128 mg/g DmE: 0.016 mg/g	Atranorin, zeorin	Singh et al. (2011)
Ramalinaceae C. Agardh
Ramalina conduplicans Vain.	ME	NS	58.2–85.4%	NS	NS	NS	Usnic acid, salazinic acid, sekikaic acid	Vinayaka et al. (2009)
Ramalina hossei Vain.	ME	NS	500 μg/ml: 89.8% + honey: 86.6%	NS	NS	NS	Usnic acid, sekikaic acid	Prashith Kekuda et al. (2009)
Ramalina pollinaria (Westr.) Ach.	ME	26.0%	No activity	NS	NS	1.0 % (w/w)	NS	Güllüce et al. (2006)
Ramalina polymorpha (Lilj.) Ach.	ME	19.0%	No activity	NS	NS	0.8 % (w/w)	NS	GüllGüllüce et al. (2006)
Ramalina terebrata Hook. F. & Taylor	ME-WE (90/10)	NS	TLC spray 2–3 antioxidant active spots	NS	NS	NVNR	NS	Bhattarai et al. (2008)
	ME-WE (90/10)	NS	302.4 μg/ml	NS	NS	NS	NS	Paudel et al. (2008)
Sphaerophoraceae Fr.
Sphaerophorus globosus (Huds.) Vain.	AcE, ME	NVNR	AcE: >1000 μg/ml ME: >1000 μg/ml	NVNR AcE > ME	NVNR AcE = ME	NS	NS	Luo et al. (2009)
Stereocaulaceae Chevall.
Stereocaulon alpinum Laurer ex Funck	ME-WE (90/10)	NS	TLC spray 2–4 antioxidant active spots	NS	NS	NVNR	NS	Bhattarai et al. (2008)
	ME-WE (90/10)	NS	409.3 μg/ml	NS	NS	NS	NS	Paudel et al. (2008)
	AcE, ME	NVNR	AcE: >1000 μg/ml ME: >1000 μg/ml	NVNR AcE > ME	NVNR AcE > ME	NS	NS	Luo et al. (2009)
Teloschistaceae Zahlbr.
Calopaca regalis (Vain.) Zahlbr.	ME-WE (90/10)	NS	TLC spray 2–3 antioxidant active spots	NS	NS	NVNR	NS	Bhattarai et al. (2008)
	ME-WE (90/10)	NS	251.7 μg/ml	NS	NS	NS	NS	Paudel et al. (2008)
Fulgensia fulgens (Sw.) Elenkin	ME	21.6%	NS	NS	0.136	12.4 mg GA/g	NS	Ranković et al. (2010b)
Xanthoria elegans (Link) Th. Fr.	AcE, DmE, EE, ME	NVNR	NVNR	NS	NS	AcE: 0.008 mg/g EE: 0.003 mg/g ME: 0.151 mg/g DmE: 0.036 mg/g	Xhantorina	Singh et al. (2011)
Trypetheliaceae Zenker
Laurera benguelensis Zahlbr.	ChE and benzene (BF) and methanol fractions (MF)	NS	ChE: 432.0 mg/ml BF: 645.8 mg/ml MF: 758.0 mg/ml	NS	NS	NS	Lichexanthone, secalonic acid D, norlichexanthon, parietin, emodin, teloschistin and citreorosein	Manojlovic et al. (2010)
Umbilicariaceae Chevall.
Lasallia pustulata (L.) Mérat	AcE, ME, WE	NS	AcE: 90.9% ME: 69.9% WE: 65.1%	AcE: 67.4% ME, WE: NVNR	NVNR AcE > ME > WE	AcE: 84.3 μg PE ME: 49.6 μg PE WE: 23.9 μg PE	NS	Kosanić et al. (2011)
Umbilicaria Antarctica Frey & Lamb	AcE, ME	NVNR	AcE: 121.3 μg/ml ME: >1000 μg/ml	AcE: 91.1% ME: NVNR	NVNR AcE > ME	NS	Lecanoric acid	Luo et al. (2009)
Umbilicaria aprina var. halei Nyl.	ME	NS	5 mg/ml: 34.6%	NS	NS	40.6 mg GA/g	Lecanoric acid	Buçukoglu et al. (2012)
Umbilicaria crustulosa (Ach.) Lamy	AcE	NS	60.1%	Moderate 42.0%	Low 0.050	39.6 μg PE/mg (28.1 μg RE/mg)	NS	Kosani ć et al. (2012b)
Umbilicaria cylindrica (L) Delise ex Duby	AcE, ME, WE	NS	NVNR	NVNR	NVNR	AcE: 19.1 μg PE ME: 42.5 μg PE WE: 19.1 μg PE (AcE: 12.9 μg RE, ME: 19.1 μg RE, WE: 11.1 μg RE)	NS	Kosani ć and Rankovic (2011)
	ME	NS	21.1% (5 mg/ml)	NS	NS	0.9 mg GA/g	NS	Buçukoglu et al. (2012)
	AcE	NS	39.4%	Moderate 33.9%	Low 0.035	19.1 μg PE/mg (12.9 μg RE/mg)	NS	Kosani ć et al. (2012b)
	ChE, ME	ChE: 29.3 μg/ml ME: 35.4 μg/ml	ChE: 31.3 μg/ml ME: 34.4 μg/ml	NS	NS	ME: 79.2 mg GA/g ChE: 71.3 mg GA/g	Salazinic acid, norstictic acid, methyl-β-orcinol carboxylate, ethyl haematommate, usnic acid, atranorin	Manojlovi ć et al. (2012b)
Umbilicaria esculenta (Miyoshi) Minks	ME	6 mg/ml: 92.1%	6 mg/ml: 1.3 mg/ml	NS	NS	1.5%	NS	Kim and Cho (2007)
Umbilicaria decussata (Vill.) Zahlbr.	ME	NS	5 mg/ml: 23.4%	NS	NS	3.3 mg GA/g	NS	Buçukoglu et al. (2012)
Umbilicaria hyperborea (Ach.) Hoffm.	AcE, DE, EE, ME	NVNR	NVNR	NS	NS	AcE: 0.001 mg/g DmE: 0.029 mg/g EE: 0.001 mg/g ME: 0.046 mg/g	Gyrophoric acid	Singh et al. (2011)
Umbilicaria leiocarpa DC.	ME	NS	5 mg/ml: 32.8%	NS	NS	35.9 mg GA/g	NS	Buçukoglu et al. (2012)
Umbilicaria nylanderiana (Zahlbr.) H. Magn.	ME	53.0%	400.2 μg/ml	NS	NS	3.0 % (w/w)	NS	Güllüce et al. (2006)
	ME	NS	5 mg/ml: 32.9%	NS	NS	48.3 mg GA/g	Umbilicaric acid	Buçukoglu et al. (2012)
Umbilicaria polyphylla (L.) Baumg.	AcE	NS	72.8%	Moderate 55.4%	Low 0.065	47.0 μg PE/mg (30.3 μg RE/mg)	NS	Kosanić et al. (2012b)
Umbilicaria virginis Schaerer	ME	NS	5 mg/ml: 20.3%	NS	NS	15.3 mg GA/g	Gyrophoric acid	Buçukoglu et al. (2012)
Verrucariaceae Zenker
Dermatcarpon intestiniformis (Körber) Hasse	ME, WE	ME: 5.3% WE: 9.5%	NS	NS	ME: 0.119 WE: 0.131	ME: 18.6 mg GA/g WE: 9.2 mg GA/g	NS	Odabasoglu et al. (2005)
Some macrostudies:
Macrostudy of 46 lichen species Highest activities: - Sticta nylanderiana Zahlbr. (S.n.) - Peltigera praetextata (Flörke ex Sommerf.) Zopf (P.p.)	ME	S.n.: 85.5% P.p.: 85.4%	S.n.: 90.4% P.p.: 87.8%	NS	S.n.: 1.5 P.p.: 1.0	S.n.: 156.1 μg CE/mg P.p.: 109.3 μg CE/mg	Sticta nylanderiana: lecanoric acid	Luo et al. (2010)
Macrostudy of 77 lichen species (Graphidaceae sp.)	DmE, ME	NS	NS	Screening of 77 extracts by two methods. Highest SO^·- RSA (>50%): Graphina multistriata Graphina salacinilabiata Graphis assamensis Graphis guimarana Graphis sikkimensis	NS	NS	NS	Behera et al. (2003)
Macrostudy of 85 lichen species	EE	NS	Screening of 99 extracts. Highest DPPH scavenging activities: Hypogymnia vittata Peltigera aphthosa Nephromopsis ornata Pseudoevernia furfuraceae Cladonia vulcani Peltigera elizobethae	NS	NS	NS	Peltigera apthosa: solorinine	Hara et al. (2011)

NS, not studied; NVNR, numeric value not reported; LP, lipoperoxidation; %INH, percentage of inhibition; SO^.− RSA, superoxide radical scavenging activity; GA, gallic acid; PE, pyrocatechol equivalent; RE, rutin equivalent; AA, ascorbic acid; T, tocopherol; TBA, thiobarbituric acid; CE, catechol equivalent; MDA, malodialdehyde; ChE, chloroform extract; ME, metanol extract; PE, petrol ether extract; AcE, acetone extract; DmE, dimethyl sulfoxide extract; EE, ethanol extract; WE, water extract; DE, diethyl-ether extract; BE, benzene extract; BuE, butanol extract; DcmE, dichlorometane extract; EaE, ethyl acetate extract.

Table 2. Antioxidant activities of isolated compounds from lichens.

Compound	Origin	LP inhibition (IC50 or %INH)	DPPH (IC50 or %INH)	SO RSA (IC50 or %INH)	Reducing power (A700 or IC50)	Reference
1,8-Dihydroxy-5-methoxy-3-methylxanthone	Pyrenula japonica Kurok [DE]	Not studied (NS)	Numeric Value Not Reported (NVNR)	NS	NS	Takenaka et al. (2000)
1,5,8-Trihydroxy-3-methylxanthone	Pyrenula japonica Kurok [DE]	NS	NVNR	NS	NS	Takenaka et al. (2000)
1,7-Dihydroxy-3-methylxanthone	Pyrenula japonica Kurok [DE]	NS	NVNR	NS	NS	Takenaka et al. (2000)
1,2,8-Trihydroxy-5-methoxy-3-methylxanthone	Pyrenula japonica Kurok [DE]	NS	NVNR	NS	NS	Takenaka et al. (2000)
1-Chloropannarin	Erioderma chilense Mont.	30% In rat brain homogenate: 27–66%	NS	NS	NS	Hidalgo et al. (1994)
7-Chlorocanarione	Lethariella cashmeriana Krog [ME]	NS	NS	Activity: 142.7	NS	Kinoshita et al. (2010)
2-O-Methylsekikaic acid	Ramalina asahinae W.L. Culb & C.F. Culb. [ME, AcE]	NS	1/IC50 = 0.052	NS	NS	Valencia-Islas et al. (2007)
Atranol	Lethariella canariensis (Ach.) Krog [ME]	NVNR	NS	NS	NS	Toledo-Marante et al. (2003)
Atranorin	Placopsis sp.	6.5% In rat brain homogenate: 7.3–9.5%	NS	NS	NS	Hidalgo et al. (1994)
	Parmotrema stuppeum (Taylor) Hale [BE, AcE]	14.0%	NS	NS	NS	Jayaprakasha et al (2000)
	Lethariella canariensis (Ach.) Krog [ME]	Unable to prevent lipid peroxidation	NS	NS	NS	Toledo-Marante et al. (2003)
	Usnea articulata (L.) Hoffm. [various extracts]	NS	NVNR	NVNR	NS	Lohézic-Le Dévéhat et al. (2007)
	Parmotrema stuppeum (Taylor) Hale [ME, AcE]	NS	1/IC50 = 0.576	NS	NS	Valencia-Islas et al. (2007)
	Parmotrema grayana Hue. [DcmE], Heterodermia obscurata (Nyl.) Trevisan [DcmE], and Cladonia sp. [DcmE]	NS	No activity	1.6%	NS	Thadhani et al. (2011)
Barbatic acid	Usnea articulata (L.) Hoffm. [various extracts]	NS	NVNR	NVNR	NS	Lohézic-Le Dévéhat et al. (2007)
Boninic acid	Ramalina asahinae W.L. Culb & C.F. Culb. [ME, AcE]	NS	1/IC50 = 0.014	NS	NS	Valencia-Islas et al. (2007)
Canarione	Lethariella sernanderi (Motyka) Obermayer [AcE]	NS	NS	Activity: 137.2	NS	Kinoshita et al. (2010)
Chloroatranol	Lethariella canariensis (Ach.) Krog [ME]	50.0–60.0%	NS	NS	NS	Toledo-Marante et al. (2003)
Chloroatranorin	Lethariella canariensis (Ach.) Krog [ME]	50.0–57.0%	NS	NS	NS	Toledo-Marante et al. (2003)
	Parmotrema stuppeum (Taylor) Hale [ME, AcE]	NS	1/IC50 = 0.589	NS	NS	Valencia-Islas et al. (2007)
Chlorohematommic acid	Lethariella canariensis (Ach.) Krog [ME]	NVNR	NS	NS	NS	Toledo-Marante et al. (2003)
Chlororubrocashmeriquinone	Lethariella sernanderi (Motyka) Obermayer [AcE], L. sinensis J.C.Wei & Y.M.Jiang [ME], and L. cashmeriana Krog [ME]	NS	NS	Activity: 277.4	NS	Kinoshita et al. (2010)
Cryptostictinolide	Usnea articulata (L.) Hoffm. [various extracts]	NS	18.0%	NVNR	NS	Lohézic-Le Dévéhat et al. (2007)
Diffractaic acid	Usnea longissima Ach. [EE]	NVNR – no antioxidant activity	No scavenging activity	NS	NS	Atalay et al. (2011)
	Protousnea magellanica (Mont.) Krog	NS	No scavenging activity	NS	NS	Brisdelli et al. (2013)
Divaricatic acid	Protousnea malacea (Stirt.) Krog	8.6% In rat brain homogenate: 1.7–8.4%	NS	NS	NS	Hidalgo et al. (1994)
	Parmotrema grayana Hue [DcmE]	NS	27.0%	92.5%	NS	Thadhani et al. (2011)
Ergosterol peroxide	Usnea articulata (L.) Hoffm. [various extracts]	NS	NVNR	NDR	NS	Lohézic-Le Dévéhat et al. (2007)
	Lobaria pulmonaria L. (Hoffm.) [AcE]	LNVNR	NVNR	NS	NS	Atalay et al. (2011)
Erythrin	Roccella montagnei Bel. [AcE]	NS	No scavenging activity	84.1%	NS	Thadhani et al. (2011)
Ethyl chlorohematommate	Lethariella canariensis (Ach.) Krog [ME]	50.0–60.0%	NS	NS	NS	Toledo-Marante et al. (2003)
Ethyl hematommate	Lethariella canariensis (Ach.) Krog [ME]	NVNR	NS	NS	NS	Toledo-Marante et al. (2003)
Evernic acid	Evernia prunastri Ach. [AcE]	NS	322.4 μg/ml	561.8 μg/ml	500 μg/ml: 0.091	Kosani ć et al. (2013)
Fumarprotocetraric acid	Usnea articulata (L.) Hoffm. [various extracts]	NS	20.0%	566.0 μM	NS	Lohézic-Le Dévéhat et al. (2007)
Gyrophoric acid	Umbilicaria virginis Schaerer	NS	50.9%	NS	NS	Buçukoglu et al. (2012)
Hematommic acid	Lethariella canariensis (Ach.) Krog [ME]	50.0–60.0%	NS	NS	NS	Toledo-Marante et al. (2003)
Isidiophorin	Lobaria pulmonaria L. (Hoffm.) [AcE]	NVNR	NVNR – high scavenging activity	NS	NS	Atalay et al. (2011)
Lecanoric acid	Parmotrema stuppeum (Taylor) Hale [BE, AcE]	12.0–36.0%	NS	NS	NS	Jayaprakasha et al (2000)
	Parmotrema tinctorum (Nyl.) Hale	NS	42.87 mM	NS	NS	Lopes et al. (2008)
Lecanoric acid	Sticta nylanderiana Zahlbr. [ME]	51.5%	35.4%	NS	NS	Luo et al. (2010)
	Parmotrema grayana Hue. [ME]	NS	34.0%	98.4%	NS	Thadhani et al. (2011)
	Umbilicaria aprina var. halei Nyl. [ME]	NS	32.5%	NS	NS	Buçukoglu et al. (2012)
Lobaric acid	Cladonia sp. [ME]	NS	No scavenging activity	96.8%	NS	Thadhani et al. (2011)
	Stereocaulon alpinum Laur.	NS	No scavenging activity	NS	NS	Brisdelli et al. (2013)
Methyl chlorohematommate	Lethariella canariensis (Ach.) Krog [ME]	66.0–70.0%	NS	NS	NS	Toledo-Marante et al. (2003)
Methyl hematommate	Lethariella canariensis (Ach.) Krog [ME]	NVNR	NS	NS	NS	Toledo-Marante et al. (2003)
	Parmotrema grayana Hue. [DcmE], and Heterodermia obscurata (Nyl.) Trevisan [DcmE]	NS	No scavenging activity	20.4%	NS	Thadhani et al. (2011)
Methyl orcinol carboxylate	Usnea articulata (L.) Hoffm. [various extracts]	NS	NVNR	NVNR	NS	Lohézic-Le Dévéhat et al. (2007)
	Heterodermia obscurata (Nyl.) Trevisan [DcmE], and Cladonia sp. [DcmE]	NS	No scavenging activity	1.5%	NS	Thadhani et al. (2011)
Methyl orsellinate	Parmotrema stuppeum (Taylor) Hale [BE, AcE]	18.0–40.0%	NS	NS	NS	Jayaprakasha et al (2000)
Methyl orsellinate	Lethariella canariensis (Ach.) Krog [ME]	NVNR	NS	NS	NS	Toledo-Marante et al. (2003)
	Parmotrema grayana Hue. [ME], Heterodermia obscurata (Nyl.) Trevisan [DcmE], and Cladonia sp. [ME]	NS	2.4%	21.2%	NS	Thadhani et al. (2011)
Montagnetol	Roccella montagnei Bél. [AcE]	NS	No scavenging activity	9.9%	NS	Thadhani et al. (2011)
Norstictic acid	Usnea articulata (L.) Hoffm. [various extracts]	NS	NVNR	580.0 μM	NS	Lohézic-Le Dévéhat et al. (2007)
	Toninia candida (Weber) Th. Fr. [AcE]	NS	102.7 μg/ml	133.5 μg/ml	500 μg/ml: 0.547	Rankovic et al. (2012)
Orcinol	Parmotrema grayana Hue. [ME]	NS	49.7%	38.2%	NS	Thadhani et al. (2011)
Orsellinic acid	Parmotrema stuppeum (Taylor) Hale. [BE, AcE]	26–50%	NS	NS	NS	Jayaprakasha et al (2000)
	Parmotrema grayana Hue. [ME]	NS	39.7%	7.9%	NS	Thadhani et al. (2011)
Pannarin	Erioderma chilense Mont.	23.0% In rat brain homogenate: 13.0–36.0%	NS	NS	NS	Hidalgo et al. (1994)
Physodic acid	Pseudoevernia furfuraceae (L.) Zopf.[AcE]	NS	69.1 μg/ml	118.2 μg/ml	500 μg/ml: 0.664	Kosanić et al. (2013)
Protolichesterinic acid	Cornicularia aculeata (Schreb.) Ach.	NS	No scavenging activity	NS	NS	Brisdelli et al. (2013)
Protrocetraric acid	Parmelia caperata (L.) Ach. [AcE]	NS	138.2 μg/ml	177.6 μg/ml	500 μg/ml: 0.090	Manojlovic et al. (2012a)
Psoromic acid	Usnea complanata (Müll. Arg.) Motyka [total extract]	0.2 mg/ml	0.271 mg/ml	NS	NS	Behera et al. (2012)
Pulmonarianin	Lobaria pulmonaria L. (Hoffm.) [AcE]	NVNR	NVNR – low scavenging activity	NS	NS	Atalay et al. (2011)
Ramalin	Ramalina terebrata Hook. f. & Taylor. [ME]	NS	0.990 μg/ml	10.2 μg/ml	0.4 μg of BHT equivalent	Paudel et al. (2011)
Rhizonaldehyde	Lobaria pulmonaria L. (Hoffm.) [AcE]	NVNR	NVNR	NS	NS	Atalay et al. (2011)
Rhizonyl alcohol	Lobaria pulmonaria L. (Hoffm.) [AcE]	NVNR	NVNR	NS	NS	Atalay et al. (2011)
Rubrocashmeriquinone	L. sernanderi (Motyka) Obermayer [AcE], and L. sinensis Wei & Jiang. [ME]	NS	NS	Activity: 87.2	NS	Kinoshita et al. (2010)
Salazinic acid	Parmelia sp. (P. saxatilis (L.) Ach.and P. sulcata Taylor.) [AcE]	NS	91.6 μg/ml	119.1 μg/ml	500 μg/ml: 0.184	Manojlovic et al. (2012a)
	Parmotrema stuppeum (Taylor) Hale. [ME, AcE]	NS	1/IC50 = 0.014	NS	NS	Valencia-Islas et al. (2007)
Sekikaic acid	Heterodermia obscurata (Nyl.) Trevis. [ME]	NS	32.6%	98.2%	NS	Thadhani et al. (2011)
Solorinine	Peltigera aphthosa (L.) Willd. [EE]	NS	120.0 μmol/ml	NS	NS	Hara et al. (2011)
Stictic acid	Usnea articulata (L.) Hoffm.[various extracts]	NS	NVNR	NVNR	NS	Lohézic-Le Dévéhat et al. (2007)
	Lobaria pulmonaria L. (Hoffm.) [AcE]	NVNR	NVNR	NS	NS	Atalay et al. (2011)
Umbilicaric acid	Umbilicaria nylanderiana (Zahlbr.) H. Magn [ME]	NS	68.1%	NS	NS	Buçukoglu et al. (2012)
Usnic acid	Lethariella canariensis (Ach.) Krog [ME]	Unable to prevent lipid peroxidation	NS	NS	NS	Toledo-Marante et al. (2003)
	Usnea articulata (L.) Hoffm.[various extracts]	NS	NVNR	NVNR	NS	Lohézic-Le Dévéhat et al. (2007)
	Ramalina asahinae W. L. Culb. & C. F. Culb. [ME, AcE]	NS	1/IC50 = 0.112	NS	NS	Valencia-Islas et al. (2007)
	Usnea longissima Ach. [EE]	NVNR – no antioxidant activity	No scavenging activity	NS	NS	Atalay et al. (2011)
	Parmotrema grayana Hue. [DcmE]	NS	No scavenging activity	59.9%	NS	Thadhani et al. (2011)
	Usnea complanata (Müll. Arg.) Motyka [Total extract]	0.214 mg/ml	0.195 mg/ml	NS	NS	Behera et al. (2012)
	Parmelia caperata (L.) Ach. [AcE]	NS	60.7 μg/ml	97.3 μg/ml	500 μg/ml: 0.547	Manojlovic et al. (2012a)
	Usnea barbata Motyka [AcE]	NS	130.7 μg/ml	197.3 μg/ml	500 μg/ml: 0.664	Rankovic et al. (2012)
	Cladonia lepidophora Ahti & Kashiw	NS	No scavenging activity	NS	NS	Brisdelli et al. (2013)
Variolaric acid	Ochrolechia deceptionis (Hue) Darb.	NS	No scavenging activity	NS	NS	Brisdelli et al. (2013)
Vesuvianic acid	Lobaria pulmonaria L. (Hoffm.) [AcE]	NVNR	NS	NS	NS	Atalay et al. (2011)
Vicanicin	Psoroma pallidum Nyl	NS	No scavenging activity	NS	NS	Brisdelli et al. (2013)
Zeorin	Cladonia sp. [DcmE]	NS	No scavenging activity	1.6%	NS	Thadhani et al. (2011)

NS, not studied; NVNR, numeric value not reported; LP, lipoperoxidation; %INH, percentage of inhibition; SO RSA, superoxide radical scavenging activity; GA, gallic acid; PE, pyrocatechol equivalent; RE, rutin equivalent; AA, ascorbic acid; T, tocopherol; TBA, thiobarbituric acid; CE, catechol equivalent; MDA, malodialdehyde; ChE, chloroform extract; ME, metanol extract; PE, petrol ether extract; AcE, acetone extract; DmE, dimethyl sulfoxide extract; EE, ethanol extract; WE, water extract; DE, diethyl-ether extract; BE, benzene extract; BuE, butanol extract; DcmE, dichlorometane extract; EaE, ethyl acetate extract.

Table 3. Evaluations of antioxidant parameters on cellular substrates.

Lichen specie/isolated compound	Solvent/origin	Cell line	LP inhibition	Antioxidant activity markers	Reference
Atranorin	Comercial	Human neuron-like cells (SH-SY5Y)	↑ Peroxyl radical-induced lipoperoxidation in vitro	Good antioxidant capacity in TRAP/TAR assays ↑ H2O2 and NO production and SO^.- scavenging activity Protection of SH-SY5Y cells against H2O2-induced cell viability impairment	Melo et al. (2011)
Cetraria islandica (L.) Ach.	Methanol	Human lymphocytes	↓ MDA levels	↑ SOD and GPx activities	Kotan et al. (2011)
Diffractaic acid	Protousnea magellanica (Mont.) Krog	HeLa cell line	Not studied (NS)	No effect on intracellular ROS level No protection against t-BHP-induced increase in intracellular ROS level	Brisdelli et al. (2013)
Evernia prunastri (L.) Ach.	Methanol	Human lymphocytes	↓ MDA levels	↑ GSH levels, SOD and GPx activities	Alpsoy et al. (2015)
Lobaric acid	Stereocaulon alpinum Laurer ex Funck	HeLa cell line	NS	No effect on intracellular ROS level No protection against t-BHP-induced increase in intracellular ROS level	Brisdelli et al. (2013)
Peltigera horizontalis (Hudson) Baumg.	Methanol	Human lymphocytes	↓ MDA levels	↑ GSH levels, SOD and GPx activities	Nardemir et al. (2013)
Peltigera praetextata (Flörke ex Sommerf.) Zopf	Methanol	Human lymphocytes	↓ MDA levels	↑ GSH levels, SOD and GPx activities	Nardemir et al. (2013)
Protolichesterinic acid	Cornicularia aculeata (Schreb.) Ach.	HeLa cell line	NS	No effect on intracellular ROS level No protection against t-BHP-induced increase in intracellular ROS level	Brisdelli et al. (2013)
Ramalin	Ramalina terebrata Hook. f. & Taylor. [ME]	Murine macrophage Raw 264.7 cells	NS	↓ NO release and H2O2 production after LPS stimulation	Paudel et al. (2011)
Salazinic acid	Xanthoparmelia cantschadalis (Ach.) Hale [ME]	Astrocyte cell line U373-MG	NS	↑ Cell viability in H2O2-treated cells ↓ ROS production ORAC value: 2.7 μmol TE/mg	de Paz et al. (2010)
Stictic acid	Xanthoparmelia conspersa (Ach.) Hale [ME]	Astrocyte cell line U373-MG	NS	↑ Cell viability in H2O2-treated cells ↓ ROS production ORAC value: 2.3 μmol TE/mg	de Paz et al. (2010)
Umbilicaria vellea (L.) Ach.	Methanol	Human lymphocytes	↓ MDA levels	↑ SOD and GPx activities (doses: 5 and 10 mg/ml)	Aslan et al. (2011)
Usnic acid (UA)	Xanthoparmelia conspersa (Ach.) Hale and X. cantschadalis (Ach.) Hale [ME]	Astrocyte cell line U373 MG	NS	↑ Cell viability in H2O2-treated cells ↓ ROS production ORAC value: 2.9 μmol TE/mg	de Paz et al. (2010)
	Comercial	Human neuron-like cells (SH-SY5Y)	↑ Lipoperoxidation	Good antioxidant capacity in TRAP/TAR assays ↓ OH^· and NO formation No effect on CAT and SOD-like activities No protection against H2O2-induced cell death ↑ ROS production	Rabelo et al. (2012)
Usnic acid (UA)	Cladonia lepidophora Ahti & Kashiw.	HeLa cell line	NS	No effect on intracellular ROS level No protection against t-BHP-induced increase in intracellular ROS level	Brisdelli et al. (2013)
Variolaric acid	Ochrolechia deceptionis (Hue) Darb.	HeLa cell line	NS	No effect on intracellular ROS level No protection against t-BHP-induced increase in intracellular ROS level	Brisdelli et al. (2013)
Vicanicin	Psoroma pallidum Nyl.	HeLa cell line	NS	No effect on intracellular ROS level No protection against t-BHP-induced increase in intracellular ROS level	Brisdelli et al. (2013)
Xantho somloensis (Gyelnik) Hale.	Methanol	Human lymphocytes	↓ MDA levels	↑ SOD and GPx activities (doses: 5 and 10 mg/ml)	Aslan et al. (2011)
Xanthoparmelia cantschadalis (Ach.) Hale	Methanol	Astrocyte cell line U373 MG	NS	↑ Cell viability in H2O2-treated cells ↓ ROS production ORAC value: 4.9 μmol TE/mg	de Paz et al. (2010)
Xanthoparmelia conspersa (Ach.) Hale	Methanol	Astrocyte cell line U373 MG	NS	↑ Cell viability in H2O2-treated cells ↓ ROS production ORAC value: 8.8 μmol TE/mg	de Paz et al. (2010)
Xanthoria elegans (Link) Th. Fr.	Water	Human lymphocytes	NS	↑ TAC level without changing TOS level	Turkez et al. (2011)

TBARS, thiobarbituric acid reactive substances; SOD, superoxide dismutase; GPx, gluthathione peroxidase; GSH, reduced gluthathione; AFB₁, aflatoxin B₁; AE, water extract; ME, methanol extract; ROS, reactive oxygen species; OH^·, hydroxyl radical; CAT, catalase; TAC, total antioxidant capacity; TOS, total oxidative stress; MPx, myeloperoxidase; cNOS, constitutive nitric oxide synthase; iNOS, inducible nitric oxide synthase; GR, gluthathione reductase; GST, glutathione transferase; DE, diethyl ether extract; AcE, acetone extract.

Table 4. In vivo evaluations of antioxidant activities.

Lichen specie/isolated compound	Solvent/origin	LP inhibition	Antioxidant enzymes and other antioxidant markers	Reference
Cetraria islandica (L.) Ach.	–	↓ MDA levels	↑ GSH levels ↓ CAT and GPx activities Association of magnesium augmented the antioxidant effect	Cernescu et al. (2011)
Diffractaic acid	Usnea longissima Ach. [DE]	↓ lipoperoxidation level in tissues	↑ SOD and GPx activities and GSH levels ↓ CAT and MPx activities ↑ cNOS activity and ↓ iNOS activity	Bayir et al. (2006)
Fumarprotocetraric acid	Cladonia verticillaris (Raddi) Fr. [AcE]	↓ endotoxin-induced lipid peroxidation	Not studied (NS)	de Barros Alves et al. (2014)
Lobaria pulmonaria (L.) Hoffm.	Methanol	↓ lipoperoxidation level in tissues	↑ SOD, GPx and GSH levels in tissues No effect on CAT and MPx levels	Karakus et al. (2009)
Peltigera rufescens (Weiss) Humb.	Methanol	↓ lipoperoxidation	↑ SOD, CAT, GSH, GR and GPx levels ↓ MPx and iNOS activities	Tanas et al. (2010)
Usnea ghattensis G. Awasthi	Methanol	↓ MDA formation in liver tissue	Depletion in antioxidant enzymes (SOD, CAT, GPx) and GSH	Verma et al. (2008a)
Usnea longissima Ach.	Water	47.1% inhibition	↑ SOD and GST levels ↓ CAT activity. Reducing power (A700): 0.1; phenolic content: 18.3 mg GA/g	Halici et al. (2005)
Usnic acid	Usnea longissima Ach. [DE]	↓ lipoperoxidation level in tissues	↑ SOD, GSH and GPx levels ↓ CAT, GR and MPx activities ↑cNOS activity and ↓iNOS activity	Odabasoglu et al. (2006)

TBARS, thiobarbituric acid reactive substances; SOD, superoxide dismutase; GPx, gluthathione peroxidase; GSH, reduced gluthathione; AFB₁, aflatoxin B₁; AE, water extract; ME, methanol extract; ROS, reactive oxygen species; OH^·, hydroxyl radical; CAT, catalase; TAC, total antioxidant capacity; TOS, total oxidative stress; MPx, myeloperoxidase; cNOS, constitutive nitric oxide synthase; iNOS, inducible nitric oxide synthase; GR, gluthathione reductase; GST, glutathione transferase; DE, diethyl ether extract; AcE, acetone extract.

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