oxidative stress: a quick primer - oxygen...

The idea is that intense training produces more free radicals than moderate exercise, which

may ―overwhelm antioxidant defenses‖ and cause ―irreparable oxidative damage… potentially resulting in ill health and/or disease,‖ writes Dr. Kelsey Fisher-Wellman, the

author of a review published in 2009.(4)

Before we dive into the evidence, let’s examine what oxidative stress is.

Oxidative Stress: A Quick Primer

During normal metabolism, your body produces unstable molecules, free radicals being the

most common. These molecules can damage your cells and create more free radicals, causing

more damage.

Your body uses different antioxidants to control free radical (oxidative) damage. The two

main kinds of antioxidants are endogenous (produced inside your body) and exogenous

(consumed from diet and other sources).

When free radicals overwhelm your antioxidant defenses, your cells are damaged. This

damage is called oxidative stress.(5,6)

There’s still some question as to whether or not oxidative stress accelerates aging.(7-

11) However, most data indicate that excessive and chronic oxidative stress may cause

cellular damage and contribute to atherosclerosis, heart disease, cancer, dementia, and

many other diseases.(12-18)

Why Oxidative Stress Isn’t Always Bad

When we first discovered that free radicals existed in humans, they were quickly blamed for

aging and other diseases.(19)

Contrary to what researchers first thought, we now know that oxidative stress is beneficial in

small amounts. In fact, it’s essential. Newer research has shown that oxidative stress prompts your cells to become stronger over time by increasing your body’s antioxidants.(20)

Free radicals also serve as important signaling molecules for a number of functions in your

body, so getting rid of them entirely would likely be counterproductive.(21)

All forms of exercise cause some oxidative stress.(22)

This is probably one of the reasons it

makes you healthier. Your body is slightly weakened, recovers, and becomes more resistant

to oxidative stress from the next workout (or other stressors).(23-30)

Most researchers agree that moderate exercise produces healthy amounts of oxidative stress,

to the point they call it an antioxidant.(31)

The concern is that doing more than that may have the opposite effect.

When Exercise May Be too Much of a Good Thing

Your body has a limited capacity to increase its antioxidants and control free radicals.

http://goo.gl/QMo47

http://www.scientificamerican.com/article.cfm?id=is-free-radical-theory-of-aging-dead

Some researchers believe that long, intense exercise (especially endurance training) may

cause more oxidative stress than humans can handle.(32)

―Heavy and sustained exercise training generates large quantities of free-radicals that likely

outstrip the buffering capacity of the system, leaving these individuals susceptible to

oxidative stress…‖ writes Harshal R. Patil, the author of a recent review on this topic.(33)

This is thought to be true ―even in superbly trained individuals.‖(34)

In theory, your body runs out of antioxidants during extreme exercise. Free radicals

overwhelm your cells and oxidative stress rises far beyond healthy levels. Over time, this

damage increases your risk of heart disease, cancer, and an early death.

Dr. James O’Keefe, the author of a review on how endurance exercise affects heart health,

states it this way:(35)

―If we went out for a run right now and you ran hard… by 60 minutes something starts happening… the free radicals blossom, and it starts burning the heart. It starts searing and

inflaming the inside of your coronary arteries.‖(36)

Every athlete has a limit to how much oxidative stress they can handle. The question is

whether or not extreme exercise can push your body past this point.

Before we look at the research, let’s define ―extreme‖ exercise.

What is “Extreme” Exercise?

If you ask most researchers in this field, the answer will probably be ―whatever causes lots of oxidative damage.‖

The problem with with this definition is that any kind of exercise can cause oxidative

damage.(37)

If your training is challenging, it’s probably causing significant oxidative stress, regardless of your sport.

(38)

This is true for:

Strength training.(39-42)

Sprinting.(43)

Motocross.(44)

Indoor rock climbing.(45)

Running.(46-49)

Cycling.(50-53)

Swimming.(54)

Rowing.(55,56)

Kayaking.(57,58)

Canoeing.(59)

Soccer.(60,61)

Tennis.(62)

Football.(63,64)

Rugby.(65,66)

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http://goo.gl/Q5MmQ

Handball.(67)

Martial arts.(68,69)

Volleyball.(70,71)

Some limited data also indicates that oxidative stress may contribute to fatigue. If this is the

case, it’s possible that any workout that can make you tired will probably cause some oxidative damage.

(72)

Is Endurance Exercise Especially Bad?

Endurance sports often get the most blame for being ―extreme‖ or ―excessive‖ when it comes to oxidative stress.

Mitochondria (the ―power plants‖ of cells) are believed to be one of the main sources of free radicals. Endurance sports use more oxygen, which sends more energy through your

mitochondria. In theory, the higher your metabolic rate and energy needs, the more free

radicals your mitochondria will produce.(73,74)

However, free radicals are also created through other pathways, and are not always related to

your oxygen needs.(75)

Several studies have shown that despite much higher oxygen intakes

during aerobic exercise, anaerobic exercise (sprinting, weight lifting, etc) can produce similar

levels of oxidative damage.(76,77)

Studies that didn’t measure oxygen consumption have also found similar levels of oxidative damage after strength and endurance training.

(78,79)

That said, it’s generally possible to do more endurance training, which will probably create more free radicals over time. No athlete runs sprints for six hours straight, but many pro

cyclists ride their bikes that long every day.

On the other hand, there are a lot of bodybuilders, football players, and other physique or

mixed sport athletes who train several hours a day.

For the rest of the article, we’ll loosely define ―extreme‖ exercise as anything that has you working at a moderate to high intensity for more than about 1-2 hours per day, endurance or

not, endurance or otherwise.

Now let’s see if this kind of training is bad for you.

What the Evidence Says About Extreme Exercise and Oxidative Stress

Despite many inconsistencies and problems with the research, most data indicate that hard

exercise increases oxidative stress.(80-84)

However, long-term studies, as opposed to short-term, have shown that extreme

training can increase your body’s antioxidant levels which can help prevent and repair

oxidative damage.(85-88)

According to Dr. Karl-Heinz Wagner, ―during exercise… ROS [free radicals] formation can stimulate adaptive mechanisms… which can lead to decreased oxidative damage or apoptosis [cell death],‖ in a review published in 2011.

(89)

In one study, untrained people ran five times per week, for an hour at a time, at 80% of their

maximum heart rates (basically, the definition of ―chronic cardio‖). There was an increase in lipid peroxidation and oxidative stress at the beginning of the study. Twelve weeks later,

however, the runners had less lipid peroxidation, higher antioxidant levels, and less oxidative

stress than before the study.(90)

Most people wouldn’t call five hours per week of running ―extreme,‖ but other studies have shown the same adaptations can occur with much more training.

With consistent training, the ―capacity of the body will expand or adapt; ultimately leading to

improvements in health and/or human performance,‖ continues Dr. Wagner.

Endurance athletes are able to prevent or at least minimize oxidative stress after races and

brutal training periods. The rise in oxidative stress is moderate to nonexistent, and usually

goes back to normal hours or days later.(91-103)

Oxidative damage sometimes occurs, but it’s minimal.(104) It’s also repaired quickly and most

data indicate there is no lasting harm.(105,106)

Multiple studies have shown that after Ironman triathlons, well-trained athletes have a large

decline in DNA damage for about three weeks.(107-109)

The researchers believe this is thanks

to ―the upregulation of repair mechanisms and enhanced endogenous antioxidative systems.‖(110)

In other words, their training increased their body’s ability to prevent and repair DNA damage, largely by increasing its antioxidant defenses.

Figure 1. DNA damage tends to decline after an Ironman in well-trained athletes. This is

because training increases your body’s antioxidants and DNA repair mechanisms.

http://goo.gl/1YlBW

http://www.marksdailyapple.com/case-against-cardio/

http://cebp.aacrjournals.org/content/17/8/1913.full.pdf

http://evidencemag.com/exercise-oxidative-stress/screen-shot-2013-01-16-at-10-51-24-am/

It’s also likely this boost in antioxidant defenses protects against oxidative stress from “non-exercise related conditions.”(111)

That is, if you get stuck in an elevator with someone reeking of cigarettes, your body will

probably suffer less damage than if you weren’t training like nuts.

Animal studies have also shown that exhaustive and moderate exercise helps increase

antioxidant levels and protect against oxidative stress.(112-117)

Elite endurance athletes (who train more than almost anyone) also tend to live longer than the

average population, although it’s impossible to tell if this is from better antioxidant protection or other reasons.

(118-120)

That said, at least one study has found higher levels of oxidative damage in well-trained

endurance athletes at rest.(121)

This is misleading, however, because the athletes had done a hard race two days before being

tested, and it’s possible this wasn’t enough time for them to recover. Even so, they were

better protected against some markers of oxidative stress after exercise than healthy non-

athletes.

Most other studies have also shown similar or lower levels of oxidative stress in athletes

compared to healthy non-athletes.(122,123)

Before you decide to do an Ironman or become Mr. Olympia however, understand that

boosting your antioxidant levels often requires a lot of training.

How Much Should You Train to Protect Your Body?

This depends on what you’re training for.

If you exercise only for health, you don’t need to train three hours a day to be well protected against oxidative stress.

Moderate endurance and strength training can significantly reduce oxidative stress and

increase your antioxidant defenses.(124-131)

It’s a different story if you’re a competitive athlete. You probably need to train more to protect yourself.

A low-volume training plan might be fine if your event is short, but it’s probably not best for longer or more intense competitions. It takes a lot of hard training to increase your

antioxidant levels enough to protect you from something like an Ironman or marathon.(132)

―The training programme must be sufficiently long and intense to trigger a consequent adaptive response of the antioxidant system and a decrease of oxidative stress,‖ writes Dr. Julien Finaud, the author of a 2006 review.

(133)

If you don’t train adequately, your antioxidant system may not be strong enough to protect you from permanent oxidative damage if you suddenly increase your training

volume. In other words, being a “weekend warrior” may not be ideal.(134)

One study found a large increase in oxidative damage after a football game. The the

researchers concluded that ―higher amounts of physical activity may be required‖ to protect

them from this kind of exercise.(135)

Athletes with the highest training volumes, experience, and fitness often have higher

antioxidant levels and lower levels of oxidative stress.(136-138)

Well-trained cyclists are also

better able to minimize oxidative stress after a workout than amateurs.(139)

Small amounts of endurance training can also increase your Vo2max before it significantly

raises your antioxidant levels.(140)

Basically, you can get fitter without necessarily being

better protected against oxidative stress.

In one of the studies mentioned previously, Ironman athletes were well protected against

oxidative stress after training 13 hours per week.(141)

Other data have shown that triathletes

training 14-17 hours per week were able to manage the oxidative stress from their

workouts.(142,143)

That’s a lot of training, but minimal considering it took these athletes around 10-12 hours to

finish an Ironman.

If your training is consistent and specific to your sport, it will probably increase your

antioxidant levels enough to protect you from excessive oxidative damage.

Of course, you can still overdo your training.

How Much is Too Much?

Some authors have suggested that there is an ―optimal level of exercise‖ and that extreme

training ―exceeds the currently undefined optimal level.‖(144)

The problem with this notion of an “optimal level of exercise” is that it’s relative to the individual and changes over time.

If an elite Ironman triathlete were to to do a two hour bike ride, they’d probably experience almost no oxidative stress. For them, that’s an easy workout.

If a couch potato who hasn’t exercised in 30 years were to do the same workout, it might cause far more damage.

Oxidative stress is not necessarily dependent on how much you exercise, but how much

you do relative to your current ability level.

In untrained people, intense, moderate, and light exercise all cause about the same amount of

oxidative damage.(145)

http://goo.gl/YYqbC

Figure 2. Intense, moderate, and easy training all increase MDA levels (a marker of oxidative

damage) about the same amount in untrained people.

According Dr. Niels Vollaard, the author of a review published in 2005, oxidative stress is

―desired, or even a required consequence of exercise, which is controlled to such an extent that it occurs whenever the relative exercise intensity is sufficiently high, regardless of

training status.‖(146)

For example, professional cyclists often train over 1,000 hours per year — over 3 hours per

day, and at times, 30-40 hours per week. Considering the demands of their sport, this might

be ―optimal.‖

They tend to have higher antioxidant levels than amateur cyclists and healthy non-athletes.

Their antioxidant levels are also increased at the end of a 20 day stage race, suggesting they

handled the oxidative stress well.(147)

Other studies have also found that well-trained cyclists are able to control the oxidative

stress from intense training and racing.(148-151)

If you push yourself outside of your comfort zone, you’re probably going to experience some oxidative stress. If you do too much, you might hurt yourself. If you don’t do enough, you may not be protected from higher training loads and other sources of oxidative stress.

This process ―can be seen as no different from other responses to exercise: a certain load disturbs homeostasis, resulting in adaptations in the body to be able to cope with a similar

load in the future,‖ writes Dr. Vollaard.(152)

Some athletes beat themselves into the ground with too much volume, intensity, or both. In

these cases, it’s possible oxidative stress may play a role in overtraining.(153,159)

Luckily, it’s usually not hard to tell when this is happening. Overtraining makes you sick,

anxious, and moody. It destroys your sleep, performance, appetite, and libido. It also

perpetuates sore or fatigued muscles, causes injuries, and usually makes you feel horrible.(160-

163)

If that sounds familiar, you’re doing too much.

The Truth about Extreme Exercise, Oxidative Stress, and Your Health

http://evidencemag.com/exercise-oxidative-stress/screen-shot-2013-01-18-at-11-50-21-am/

http://goo.gl/FbLpa

http://goo.gl/hTQaS

Every kind of hard exercise creates free radicals that cause some oxidative damage.

This damage is quickly repaired, and sparks adaptations that make you more resistant to

oxidative damage from high volume and/or intensity training.

With more training, your body increases its antioxidant levels to control the additional free

radicals, generally keeping oxidative stress within safe limits.

It’s possible to hurt yourself by overtraining, and oxidative stress may be part of this process.

However, it’s not news that overtraining is bad for you. High volume training is also not

the same thing as overtraining.

If you train smart by providing enough stimulus to progress, but not so much that you

overtrain, the oxidative stress from your training will probably not damage your health.

At present, ―there are no indications that exercise-induced oxidative stress has any negative

impact on health…,‖ according to Dr. Vollaard.(164)

As with fatigue, muscle soreness, and any other response to exercise, oxidative stress should

not be avoided, but managed with smart, consistent training.

You don’t need to do extreme athletics to be healthy.

On the other hand, if you do train hours a day for extreme sports, you probably don’t have to worry about it damaging your health or shortening your life. It might even make you

healthier.

Most evidence indicates that if you train in a progressive, intelligent manner, with

adequate recovery between workouts, you can build up to extremely high training loads

and still be protected against potentially dangerous levels of oxidative stress.

You’re reading Part 5 of a series on whether or not exercise damages your heart. Click here

to read Part 6.

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exercise. Free Radic Res. 2005 Dec;39(12):1317-24. Abstract: http://pmid.us/16298861 | Full Text: NA

52 Tauler P, Sureda A, Cases N, et al. Increased lymphocyte antioxidant defences in response to exhaustive exercise do not prevent oxidative

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53 Martarelli D, Pompei P. Oxidative stress and antioxidant changes during a 24-hours mountain bike endurance exercise in master athletes.

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54 Deminice R, Trindade CS, Degiovanni GC, et al. Oxidative stress biomarkers response to high intensity interval training and relation to

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NA

55 Kyparos A, Riganas C, Nikolaidis MG, et al. The effect of exercise-induced hypoxemia on blood redox status in well-trained rowers. Eur

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56 Kyparos A, Vrabas IS, Nikolaidis MG, et al. Increased oxidative stress blood markers in well-trained rowers following two thousand-

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Full Text: NA

59 Teixeira V, Valente H, Casal S, et al. Antioxidant status, oxidative stress, and damage in elite trained kayakers and canoeists and

sedentary controls. Int J Sport Nutr Exerc Metab. 2009 Oct;19(5):443-56. Abstract: http://pmid.us/19910648 | Full Text: NA

60 Fatouros IG, Chatzinikolaou A, Douroudos II, et al. Time-course of changes in oxidative stress and antioxidant status responses following

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62 Kyparos A, Salonikidis K, Nikolaidis MG, et al. Short duration exhaustive aerobic exercise induces oxidative stress: a novel play-oriented

volitional fatigue test. J Sports Med Phys Fitness. 2007 Dec;47(4):483-90. Abstract: http://pmid.us/18091691 | Full Text: NA

63 Rudarli Nalçakan G, Nalçakan M, Var A, et al. Acute oxidative stress and antioxidant status responses following an American football

match. J Sports Med Phys Fitness. 2011 Sep;51(3):533-9. Abstract: http://pmid.us/21904294 | Full Text: NA

64 Schippinger G, Wonisch W, Abuja PM, et al. Lipid peroxidation and antioxidant status in professional American football players during

competition. Eur J Clin Invest. 2002 Sep;32(9):686-92. Abstract: http://pmid.us/12486869 | Full Text: NA

65 Chang CK, Tseng HF, Hsuuw YD, et al. Higher LDL oxidation at rest and after a rugby game in weekend warriors. Ann Nutr Metab.

2002;46(3-4):103-7. Abstract: http://pmid.us/12169852 | Full Text: NA









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66 Finaud J, Scislowski V, Lac G, et al. Antioxidant status and oxidative stress in professional rugby players: evolution throughout a season.

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67 Marin DP, Macedo dos Santos RC, Bolin AP. Cytokines and Oxidative Stress Status Following a Handball Game in Elite Male Players.

Oxid Med Cell Longev. 2011; 2011: 804873. Full Text: http://goo.gl/yhar9

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70 Martinovic J, Dopsaj V, Dopsaj MJ, et al. Long-term effects of oxidative stress in volleyball players. Int J Sports Med. 2009

Dec;30(12):851-6. doi: 10.1055/s-0029-1238289. Abstract: http://pmid.us/20013555 | Full Text: NA

71 Martinović J, Dopsaj V, Kotur-Stevuljević J, et al. Oxidative stress biomarker monitoring in elite women volleyball athletes during a 6-

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author.



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| Full Text: http://goo.gl/bKVQT

76 Alessio HM, Hagerman AE, Fulkerson BK, et al. Generation of reactive oxygen species after exhaustive aerobic and isometric exercise.

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77 Shi M, Wang X, Yamanaka T, et al. Effects of anaerobic exercise and aerobic exercise on biomarkers of oxidative stress. Environ Health

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78 Bloomer RJ, Goldfarb AH, Wideman L, et al. Effects of acute aerobic and anaerobic exercise on blood markers of oxidative stress. J

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83 König D, Neubauer O, Nics L, et al. Biomarkers of exercise-induced myocardial stress in relation to inflammatory and oxidative stress.

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84 Clarkson PM, Thompson HS. Antioxidants: what role do they play in physical activity and health? Am J Clin Nutr 2000;72:637 – 46.

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86 Knez WL, Jenkins DG, Coombes JS. Oxidative stress in half and full Ironman triathletes. Med Sci Sports Exerc. 2007 Feb;39(2):283-8.

Abstract: http://pmid.us/17277592 | Full Text: http://goo.gl/5uqlb



http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3172986/

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90 Miyazaki H, Oh-ishi S, Ookawara T, et al. Strenuous endurance training in humans reduces oxidative stress following exhausting

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91 Wagner KH, Reichhold S, Hölzl C, et al. Well-trained, healthy triathletes experience no adverse health risks regarding oxidative stress and

DNA damage by participating in an ultra-endurance event. Toxicology. 2010 Dec 5;278(2):211-6. doi: 10.1016/j.tox.2009.09.006. Epub

2009 Sep 18. Abstract: http://pmid.us/19766696 | Full Text: Requested.

92 Aguiló A, Tauler P, Fuentespina E, et al. Antioxidant response to oxidative stress induced by exhaustive exercise. Physiol Behav. 2005

Jan 31;84(1):1-7. Epub 2004 Nov 10. Abstract: http://pmid.us/15642600 | Full Text: http://goo.gl/6Q3Rq

93 Reichhold S, Neubauer O, Veronika E, et al. No acute and persistent DNA damage after an Ironman triathlon. Cancer Epidemiol

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94 Reichhold S, Neubauer O, Hoelzl C, et al. DNA damage in response to an Ironman triathlon. Free Radic Res. 2009 Aug;43(8):753-60.

doi: 10.1080/10715760903040628. Abstract: http://pmid.us/19591014 | Full Text: NA

95 Cases N, Sureda A, Maestre I, et al. Response of antioxidant defences to oxidative stress induced by prolonged exercise: antioxidant

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Full Text: NA

96 Margaritis I, Tessier F, Richard MJ, et al. No evidence of oxidative stress after a triathlon race in highly trained competitors. Int J Sports

Med. 1997 Apr;18(3):186-90. Abstract: http://pmid.us/9187972 | Full Text: NA

97 Shing CM, Peake JM, Ahern SM, et al. The effect of consecutive days of exercise on markers of oxidative stress. Appl Physiol Nutr

Metab. 2007 Aug;32(4):677-85. Abstract: http://pmid.us/17622282 | Full Text: http://goo.gl/4CguF

98 Serrano E, Venegas C, Escames G, et al. Antioxidant defence and inflammatory response in professional road cyclists during a 4-day

competition. J Sports Sci. 2010 Aug;28(10):1047-56. Abstract: http://pmid.us/20686993 | Full Text: NA

99 Kabasakalis A, Kyparos A, Tsalis G, et al. Blood oxidative stress markers after ultramarathon swimming. J Strength Cond Res. 2011

Mar;25(3):805-11. doi: 10.1519/JSC.0b013e3181d0b109. Abstract: http://pmid.us/20613649 | Full Text: NA

100 Okamura K, Doi T, Hamada K, et al. Effect of repeated exercise on urinary 8-hydroxy-deoxyguanosine excretion in humans. Free Radic

Res. 1997 Jun;26(6):507-14. Abstract: http://pmid.us/9212344 | Full Text: NA

101 Neubauer O, Reichhold S, Nersesyan A, et al. Exercise-induced DNA damage: is there a relationship with inflammatory responses?

Exerc Immunol Rev. 2008;14:51-72. Abstract: http://pmid.us/19203084 | Full Text: http://goo.gl/s0fhs



103 Tong TK, Lin H, Lippi G, et al. Serum oxidant and antioxidant status in adolescents undergoing professional endurance sports training.

Oxid Med Cell Longev. 2012;2012:741239. doi: 10.1155/2012/741239. Epub 2012 Apr 17. Abstract: http://pmid.us/22577491 | Full Text:

http://goo.gl/5v58a



105 Neubauer O, Reichhold S, Nersesyan A, et al. Exercise-induced DNA damage: is there a relationship with inflammatory responses?

Exerc Immunol Rev. 2008;14:51-72. Abstract: http://pmid.us/19203084 | Full Text: http://goo.gl/s0fhs







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108 Reichhold S, Neubauer O, Hoelzl C, et al. DNA damage in response to an Ironman triathlon. Free Radic Res. 2009 Aug;43(8):753-60.

doi: 10.1080/10715760903040628. Abstract: http://pmid.us/19591014 | Full Text: NA

109 Wagner KH, Reichhold S, Hölzl C, et al. Well-trained, healthy triathletes experience no adverse health risks regarding oxidative stress

and DNA damage by participating in an ultra-endurance event. Toxicology. 2010 Dec 5;278(2):211-6. doi: 10.1016/j.tox.2009.09.006. Epub

2009 Sep 18. Abstract: http://pmid.us/19766696 | Full Text: Requested.





112 Oztasan N, Taysi S, Gumustekin K, et al. Endurance training attenuates exercise-induced oxidative stress in erythrocytes in rat. Eur J

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127 Radak Z, Chung HY, Goto S. Exercise and hormesis: oxidative stress-related adaptation for successful aging. Biogerontology.

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128 Radak Z, Chung HY, Goto S. Systemic adaptation to oxidative challenge induced by regular exercise. Free Radic Biol Med. 2008 Jan

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129 Majerczak J, Rychlik B, Grzelak A, et al. Effect of 5-week moderate intensity endurance training on the oxidative stress, muscle specific

uncoupling protein (UCP3) and superoxide dismutase (SOD2) contents in vastus lateralis of young, healthy men. J Physiol Pharmacol. 2010

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130 Campbell PT, Gross MD, Potter JD, et al. Effect of exercise on oxidative stress: a 12-month randomized, controlled trial. Med Sci Sports

Exerc. 2010 Aug;42(8):1448-53. doi: 10.1249/MSS.0b013e3181cfc908. Abstract: http://pmid.us/20139793 | Full Text: http://goo.gl/PcLYx

131 Fatouros IG, Jamurtas AZ, Villiotou V, et al. Oxidative stress responses in older men during endurance training and detraining. Med Sci

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132 Finaud J, Lac G, Filaire E. Oxidative stress: relationship with exercise and training. Sports Med. 2006;36(4):327-58. Abstract:

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134 Chang CK, Tseng HF, Hsuuw YD, et al. Higher LDL oxidation at rest and after a rugby game in weekend warriors. Ann Nutr Metab.

2002;46(3-4):103-7. Abstract: http://pmid.us/12169852 | Full Text: NA

135 Rudarli Nalçakan G, Nalçakan M, Var A, et al. Acute oxidative stress and antioxidant status responses following an American football

match. J Sports Med Phys Fitness. 2011 Sep;51(3):533-9. Abstract: http://pmid.us/21904294 | Full Text: NA

136 Margaritis I, Tessier F, Richard MJ, et al. No evidence of oxidative stress after a triathlon race in highly trained competitors. Int J Sports

Med. 1997 Apr;18(3):186-90. Abstract: http://pmid.us/9187972 | Full Text: NA

137 Martinovic J, Dopsaj V, Dopsaj MJ, et al. Long-term effects of oxidative stress in volleyball players. Int J Sports Med. 2009

Dec;30(12):851-6. doi: 10.1055/s-0029-1238289. Abstract: http://pmid.us/20013555 | Full Text: NA



139 Mena P, Maynar M, Gutierrez JM, et al. Erythrocyte free radical scavenger enzymes in bicycle professional racers. Adaptation to

training. Int J Sports Med. 1991 Dec;12(6):563-6. Abstract: http://pmid.us/1797698 | Full Text: NA











145 Moflehi D, Kok LY, Tengku-Kamalden TF, et al. Effect of single-session aerobic exercise with varying intensities on lipid peroxidation

and muscle-damage markers in sedentary males. Glob J Health Sci. 2012 May 23;4(4):48-54. doi: 10.5539/gjhs.v4n4p48. Abstract:

http://pmid.us/22980341 | Full Text: http://goo.gl/FbLpa



147 Mena P, Maynar M, Gutierrez JM, et al. Erythrocyte free radical scavenger enzymes in bicycle professional racers. Adaptation to

training. Int J Sports Med. 1991 Dec;12(6):563-6. Abstract: http://pmid.us/1797698 | Full Text: NA

148 Shing CM, Peake JM, Ahern SM, et al. The effect of consecutive days of exercise on markers of oxidative stress. Appl Physiol Nutr

Metab. 2007 Aug;32(4):677-85. Abstract: http://pmid.us/17622282 | Full Text: http://goo.gl/4CguF






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150 Aguiló A, Tauler P, Fuentespina E, et al. Antioxidant response to oxidative stress induced by exhaustive exercise. Physiol Behav. 2005

Jan 31;84(1):1-7. Epub 2004 Nov 10. Abstract: http://pmid.us/15642600 | Full Text: http://goo.gl/6Q3Rq

151 Tong TK, Lin H, Lippi G, et al. Serum oxidant and antioxidant status in adolescents undergoing professional endurance sports training.

Oxid Med Cell Longev. 2012;2012:741239. doi: 10.1155/2012/741239. Epub 2012 Apr 17. Abstract: http://pmid.us/22577491 | Full Text:

http://goo.gl/5v58a



153 Schippinger G, Fankhauser F, Abuja PM, et al. Competitive and seasonal oxidative stress in elite alpine ski racers. Scand J Med Sci

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2004;34(14):967-81. Abstract: http://pmid.us/15571428 | Full Text: http://goo.gl/uu8Bn



Bald, blind, never get cancer, resistant to low oxygen. Look like hotdogs with teeth. Live for

thirty years. Teeth stick out through their skin. Live in colony- air supply is very limited- use

up much of the oxygen and generated high levels of carbon dioxide. Brain is very protective

against low oxygen- taking tens of millions of years.



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Live – how is it that breathing in high levels of carbon dioxide is not painful.

Live in central east Africa where they are considered to be a pest eating the vegetables of the

farmer from underground.

Free Radic Biol Med. 2008 Jan 15;44(2):126-31. doi: 10.1016/j.freeradbiomed.2007.02.001.

Epub 2007 Feb 9.

Moderate exercise is an antioxidant:

upregulation of antioxidant genes by

training.

Gomez-Cabrera MC1, Domenech E, Viña J.

Author information

Abstract

Exercise causes oxidative stress only when exhaustive. Strenuous exercise causes oxidation

of glutathione, release of cytosolic enzymes, and other signs of cell damage. However, there

is increasing evidence that reactive oxygen species (ROS) not only are toxic but also play an

important role in cell signaling and in the regulation of gene expression. Xanthine oxidase is

involved in the generation of superoxide associated with exhaustive exercise. Allopurinol (an

inhibitor of this enzyme) prevents muscle damage after exhaustive exercise, but also modifies

cell signaling pathways associated with both moderate and exhaustive exercise in rats and

humans. In gastrocnemius muscle from rats, exercise caused an activation of MAP kinases.

This in turn activated the NF-kappaB pathway and consequently the expression of important

enzymes associated with defense against ROS (superoxide dismutase) and adaptation to

exercise (eNOS and iNOS). All these changes were abolished when ROS production was

prevented by allopurinol. Thus ROS act as signals in exercise because decreasing their

formation prevents activation of important signaling pathways that cause useful adaptations

in cells. Because these signals result in an upregulation of powerful antioxidant enzymes,

exercise itself can be considered an antioxidant. We have found that interfering with free

radical metabolism with antioxidants may hamper useful adaptations to training.

Curr Pharm Des. 2011;17(22):2290-307.


http://www.ncbi.nlm.nih.gov/pubmed?term=Gomez-Cabrera%20MC%5BAuthor%5D&cauthor=true&cauthor_uid=18191748

http://www.ncbi.nlm.nih.gov/pubmed?term=Domenech%20E%5BAuthor%5D&cauthor=true&cauthor_uid=18191748

http://www.ncbi.nlm.nih.gov/pubmed?term=Vi%C3%B1a%20J%5BAuthor%5D&cauthor=true&cauthor_uid=18191748



Walking the oxidative stress tightrope: a

perspective from the naked mole-rat, the

longest-living rodent.

Rodriguez KA1, Wywial E, Perez VI, Lambert AJ, Edrey YH, Lewis KN, Grimes K, Lindsey

ML, Brand MD, Buffenstein R.

Author information

Abstract

Reactive oxygen species (ROS), by-products of aerobic metabolism, cause oxidative damage

to cells and tissue and not surprisingly many theories have arisen to link ROS-induced

oxidative stress to aging and health. While studies clearly link ROS to a plethora of divergent

diseases, their role in aging is still debatable. Genetic knock-down manipulations of

antioxidants alter the levels of accrued oxidative damage, however, the resultant effect of

increased oxidative stress on lifespan are equivocal. Similarly the impact of elevating

antioxidant levels through transgenic manipulations yield inconsistent effects on longevity.

Furthermore, comparative data from a wide range of endotherms with disparate longevity

remain inconclusive. Many long-living species such as birds, bats and mole-rats exhibit high-

levels of oxidative damage, evident already at young ages. Clearly, neither the amount of

ROS per se nor the sensitivity in neutralizing ROS are as important as whether or not the

accrued oxidative stress leads to oxidative-damage-linked age-associated diseases. In this

review we examine the literature on ROS, its relation to disease and the lessons gleaned from

a comparative approach based upon species with widely divergent responses. We specifically

focus on the longest lived rodent, the naked mole-rat, which maintains good health and

provides novel insights into the paradox of maintaining both an extended healthspan and

lifespan despite high oxidative stress from a young age.

aked Mole-Rat Unfazed By Oxidative Stress

Date:

October 10, 2006

Source:

American Physiological Society

Summary:

The long-lived naked mole-rat shows much higher levels of oxidative stress and damage and less robust repair mechanisms

than the short-lived mouse, findings that could change the oxidative stress theory of aging. The new study comparing the

naked mole-rat, which has a life span of 28 years, and the mouse, which has a lifespan of three years, will be presented at

The American Physiological Society conference, Comparative Physiology 2006: Integrating Diversity.

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http://www.ncbi.nlm.nih.gov/pubmed?term=Rodriguez%20KA%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Wywial%20E%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Perez%20VI%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Lambert%20AJ%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Edrey%20YH%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Lewis%20KN%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Grimes%20K%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Lindsey%20ML%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Lindsey%20ML%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Brand%20MD%5BAuthor%5D&cauthor=true&cauthor_uid=21736541

http://www.ncbi.nlm.nih.gov/pubmed?term=Buffenstein%20R%5BAuthor%5D&cauthor=true&cauthor_uid=21736541


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A naked mole rat (Heterocephalus glaber).

Credit: Photo Roman Klementschitz, Wien / courtesy of Wikimedia Commons

The long-lived naked mole-rat shows much higher levels of oxidative stress and damage and less robust repair

mechanisms than the short-lived mouse, findings that could change the oxidative stress theory of aging.

The new study comparing the naked mole-rat, which has a life span of 28 years, and the mouse, which has a lifespan of three

years, will be presented Oct. 8 at The American Physiological Society conference, Comparative Physiology 2006:

Integrating Diversity. The results fly in the face of the oxidative stress theory of aging, which holds that damage caused by

oxidative stress is a significant contributor to the aging process.

Under this theory, naked mole-rats should be better at preventing or repairing oxidative stress than their much shorter-lived

cousin, the mouse. The study, "High oxidative damage levels in the longest-living rodent, the naked mole-rat," was done by

Blazej Andziak and Rochelle Buffenstein, of The City College of New York, Timothy P. O'Connor, of Weill Medical

College of Cornell University, and Asish P. Chaudhiri and Holly Van Remmen of the University of Texas Health Science

Center, San Antonio. The study was presented during a poster session on October 8.

Don't toss the oxidative stress theory of aging out the window just yet, but prepare to modify it, said Buffenstein, the senior

author. Her team suspects that the naked mole-rat's longevity stems from its ability to defend against acute bouts of oxidative

stress. That is, the kind of oxidation that happens because of an unusual occurrence rather than the kind that happens as a

result of normal aerobic respiration.

For example, when hydrogen peroxide is added to a culture containing naked mole-rat fibroblast cells, they remain viable

and appear to repair the acute damage more rapidly than shorter-lived animals, explained Buffenstein.

What is old age?

We know that all organisms age and die. It's such an inevitable course of events that most of us spend more time thinking

about how to hide the wrinkles and gray hair than we do about what our cells are actually doing to usher us to the end.

Physiologists are looking at molecules and cells to understand this process.

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One way to look at aging is to compare closely related organisms with different life spans. That's why it made sense to

compare mole-rats and mice: They're the same size and they're rodents, but the mole-rat lives to 28 years, about nine-times

longer than the mouse.

"mole-rats must have something happening at the biochemical level to allow them to do this," said Andziak, the study's lead

author. Specifically, he wanted to see if oxidative stress could explain the difference.

Oxidative stress occurs during metabolism when oxygen (O2) splits into single oxygen atoms, known as free radicals. These

oxygen atoms may circulate by themselves, or combine with other atoms and molecules to form reactive oxygen species

(ROS). ROS can damage DNA, lipids and proteins thus impairing normal cellular function. Antioxidants help to neutralize

ROS, thus restricting the potential of ROS to damage biological molecules.

mole-rat has more oxidative stress

The study compared two-year-old naked mole-rats to four-month-old mice. The researchers chose those ages so that the

animals would be equivalent ages relative to their maximum lifespans, Andziak said.

First, the researchers compared the ratio of reduced glutathione, an antioxidant, to oxidized glutathione. As the body uses up

its reduced glutathione to fight oxidative stress, the pool of oxidized glutathione increases. This ratio of reduced to oxidized

glutathione is thus an indicator of oxidative stress: the greater the ratio, the less oxidative stress has occurred. The oxidative

stress theory predicts that in naked mole-rats this ratio will be higher than in mice.

When the researchers measured this ratio in the liver, they found that the opposite was true. mole-rats had less reduced

glutathione and thus a lower ratio, indicating the mole-rat experienced much more oxidative stress. These results fit with the

findings of a previous study in which Andziak found that naked mole-rats did not have superior antioxidant capacity when

compared to mice. mole-rats had much lower activity of the ubiquitous antioxidant enzyme, cellular glutathione peroxidase.

mole-rat shows greater oxidative damage

The researchers next looked at how much damage the oxidation had caused. It is possible, they reasoned, that the mole-rat

suffers greater oxidative stress, but its physiology had somehow prevented damage from occurring.

The researchers measured oxidative damage in lipids, DNA and proteins and found that naked mole-rats showed much

greater levels of damage to each of these biological molecules, in all tissues assayed, when compared to mice. The study

found multiple signs of lipid damage: The level of isoprostanes found in the urine was 10 times higher in the naked mole-rat,

the level of malondialdehyde in liver tissue was twice as high and isoprostane levels in heart tissue was two-and-a-half times

the level of the mice.

The researchers found significantly more protein damage in the kidney and in the heart. DNA damage was greater in the

kidney and liver.

"All of the classical measures of oxidative stress are higher in the mole-rat," Andziak concluded. "Given that naked mole-

rats live an order of magnitude longer than predicted based on their body size, our findings strongly suggest that mechanisms

other than attenuated oxidative stress may explain the impressive longevity of this species."

Next steps

The next step is to determine how the mole-rats manage to live with the damage caused by oxidative stress. Buffenstein said

she suspects that the mole-rat is able to fend off the occasional oxidative insult that can occur, and that may be more

important than what happens with the steady-state levels of oxidative stress that result from normal metabolic activity.

Buffenstein theorizes that the naked mole-rats in her laboratory suffer higher levels of oxidative stress than they would in

their natural underground habitat, where they encounter much lower levels of oxygen. But this exposure at an early age may

provide some protection against acute oxidative stress and may be of considerable importance in their resistance to bursts

oxidative stressors throughout their lives, she said.

"The naked mole-rat, with its surprisingly long lifespan and remarkably delayed aging, seems like the perfect model to

provide answers about how we age and how to retard the aging process," Buffenstein said. "This animal may one day

provide the clues to how we can significantly extend life."

Gerontology. 2012;58(5):453-62. doi: 10.1159/000335966. Epub 2012 May 10.

Stress resistance in the naked mole-rat: the

bare essentials - a mini-review.

Lewis KN1, Mele J, Hornsby PJ, Buffenstein R.

Author information

Abstract

BACKGROUND:

Studies comparing similar-sized species with disparate longevity may elucidate novel

mechanisms that abrogate aging and prolong good health. We focus on the longest living

rodent, the naked mole-rat. This mouse-sized mammal lives ~8 times longer than do mice

and, despite high levels of oxidative damage evident at a young age, it is not only very

resistant to spontaneous neoplasia but also shows minimal decline in age-associated

physiological traits.

OBJECTIVES:

We assess the current status of stress resistance and longevity, focusing in particular on the

molecular and cellular responses to cytotoxins and other stressors between the short-lived

laboratory mouse and the naked mole-rat.

RESULTS:

Like other experimental animal models of lifespan extension, naked mole-rat fibroblasts are

extremely tolerant of a broad spectrum of cytotoxins including heat, heavy metals, DNA-

damaging agents and xenobiotics, showing LD(50) values between 2- and 20-fold greater

than those of fibroblasts of shorter-lived mice. Our new data reveal that naked mole-rat

fibroblasts stop proliferating even at low doses of toxin whereas those mouse fibroblasts that

survive treatment rapidly re-enter the cell cycle and may proliferate with DNA damage.



http://www.ncbi.nlm.nih.gov/pubmed?term=Mele%20J%5BAuthor%5D&cauthor=true&cauthor_uid=22572398

http://www.ncbi.nlm.nih.gov/pubmed?term=Hornsby%20PJ%5BAuthor%5D&cauthor=true&cauthor_uid=22572398



Naked mole-rat fibroblasts also show significantly higher constitutive levels of both p53 and

Nrf2 protein levels and activity, and this increases even further in response to toxins.

CONCLUSION:

Enhanced cell signaling via p53 and Nrf2 protects cells against proliferating with damage,

augments clearance of damaged proteins and organelles and facilitates the maintenance of

both genomic and protein integrity. These pathways collectively regulate a myriad of

mechanisms which may contribute to the attenuated aging profile and sustained healthspan of

the naked mole-rat. Understanding how these are regulated may be also integral to sustaining

positive human healthspan well into old age and may elucidate novel therapeutics for

delaying the onset and progression of physiological declines that characterize the aging

process.

Antioxid Redox Signal. 2013 Oct 20;19(12):1388-99. doi: 10.1089/ars.2012.4911. Epub 2012

Dec 7.

The naked mole-rat response to oxidative

stress: just deal with it.

Lewis KN1, Andziak B, Yang T, Buffenstein R.

Author information

Abstract

SIGNIFICANCE:

The oxidative stress theory of aging has been the most widely accepted theory of aging

providing insights into why we age and die for over 50 years, despite mounting evidence

from a multitude of species indicating that there is no direct relationship between reactive

oxygen species (ROS) and longevity. Here we explore how different species, including the

longest lived rodent, the naked mole-rat, have defied the most predominant aging theory.

RECENT ADVANCES:

In the case of extremely long-lived naked mole-rat, levels of ROS production are found to be

similar to mice, antioxidant defenses unexceptional, and even under constitutive conditions,

naked mole-rats combine a pro-oxidant intracellular milieu with high, steady state levels of

oxidative damage. Clearly, naked mole-rats can tolerate this level of oxidative stress and

must have mechanisms in place to prevent its translation into potentially lethal diseases.

CRITICAL ISSUES:

In addition to the naked mole-rat, other species from across the phylogenetic spectrum and

even certain mouse strains do not support this theory. Moreover, overexpressing or knocking

down antioxidant levels alters levels of oxidative damage and even cancer incidence, but does

not modulate lifespan.



http://www.ncbi.nlm.nih.gov/pubmed?term=Andziak%20B%5BAuthor%5D&cauthor=true&cauthor_uid=23025341

http://www.ncbi.nlm.nih.gov/pubmed?term=Yang%20T%5BAuthor%5D&cauthor=true&cauthor_uid=23025341



FUTURE DIRECTIONS:

Perhaps, it is not oxidative stress that modulates healthspan and longevity, but other

cytoprotective mechanisms that allow animals to deal with high levels of oxidative damage

and stress, and nevertheless live long, relatively healthy lifespans. Studying these

mechanisms in uniquely long-lived species, like the naked mole-rat, may help us tease out the

key contributors to aging and longevity.

Proc Natl Acad Sci U S A. 2009 Nov 17;106(46):19352-7. doi: 10.1073/pnas.0905252106.

Epub 2009 Oct 26.

Hypersensitivity to contact inhibition

provides a clue to cancer resistance of

naked mole-rat.

Seluanov A1, Hine C, Azpurua J, Feigenson M, Bozzella M, Mao Z, Catania KC, Gorbunova

V.

Author information

Abstract

The naked mole-rat is the longest living rodent with a maximum lifespan exceeding 28 years.

In addition to its longevity, naked mole-rats have an extraordinary resistance to cancer as

tumors have never been observed in these rodents. Furthermore, we show that a combination

of activated Ras and SV40 LT fails to induce robust anchorage-independent growth in naked

mole-rat cells, while it readily transforms mouse fibroblasts. The mechanisms responsible for

the cancer resistance of naked mole-rats were unknown. Here we show that naked mole-rat

fibroblasts display hypersensitivity to contact inhibition, a phenomenon we termed "early

contact inhibition." Contact inhibition is a key anticancer mechanism that arrests cell division

when cells reach a high density. In cell culture, naked mole-rat fibroblasts arrest at a much

lower density than those from a mouse. We demonstrate that early contact inhibition requires

the activity of p53 and pRb tumor suppressor pathways. Inactivation of both p53 and pRb

attenuates early contact inhibition. Contact inhibition in human and mouse is triggered by the

induction of p27(Kip1). In contrast, early contact inhibition in naked mole-rat is associated

with the induction of p16(Ink4a). Furthermore, we show that the roles of p16(Ink4a) and


http://www.ncbi.nlm.nih.gov/pubmed?term=Seluanov%20A%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Hine%20C%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Azpurua%20J%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Feigenson%20M%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Bozzella%20M%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Mao%20Z%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Catania%20KC%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Gorbunova%20V%5BAuthor%5D&cauthor=true&cauthor_uid=19858485

http://www.ncbi.nlm.nih.gov/pubmed?term=Gorbunova%20V%5BAuthor%5D&cauthor=true&cauthor_uid=19858485


p27(Kip1) in the control of contact inhibition became temporally separated in this species:

the early contact inhibition is controlled by p16(Ink4a), and regular contact inhibition is

controlled by p27(Kip1). We propose that the additional layer of protection conferred by two-

tiered contact inhibition contributes to the remarkable tumor resistance of the naked mole-rat.

oxidative stress: a quick primer - oxygen...

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