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WHITE PAPER by Lena Burri, PhD
THE IMPORTANCE OF CHOLINE
Choline is a conditionally essential nutrient,
meaning that it can be produced by the body,
but under certain conditions the supply might be
inadequate [1]. Dogs need choline for the synthesis of
neurotransmitters (i.e. acetylcholine) for memory and
muscle function. Choline is also part of phospholipids
(i.e. phosphatidylcholine), which are important for
transport of lipids, building of cell membranes and
cell membrane signaling [2].
compounds made from choline, transport and pro-
vide methyl groups, which are used by the body to
control protein function and gene expression im-
portant for the proper functioning of the cardiovas-
cular, neurological, reproductive and detoxification
systems.
Moreover, choline has a role in being a methyl
donor. Inasmuch, choline can be processed to
betaine, which then provides a methyl group for the
generation of methionine. Methionine is an essential
amino acid used to build proteins. It is made from
homocysteine, thereby reducing the heart disease
risk, which is associated with homocysteine. These
Many of these pathways are challenged during in-
tense physical activity of long duration, such as dog
sled races, and the demand for free, non-membrane
bound choline is increased to counteract the de-
crease of plasma choline concentrations during
endurance exercise [3, 4]. The exact reasons for why
choline concentrations decline during a high exercise
performance are unclear, but were suggested to not
necessarily reflect choline utilization, instead it might
be a consequence of redistribution of fluid pools
during exertion [5].
Figure 1: Structural function of phosphatidylcholine
Water loving head groups
Micelles Liposome
Bilayer Membrane
CHOLINE
S-adenosyl-homocysteine Phosphatidylethanolamine
PhosphatidylcholineCell membrane & lipid transport component
S-adenosyl-methionineMethyl donor
HomocysteineHeart disease risk factor
Cysteine
Methionine Dimethylglycine
Betaine Choline
AcetylcholineNeurotransmitter
Figure 2: Schematic of the conversion pathwaysof phosphatidylcholine to its metabolites
- 2 -
Hydrophopic fatty acids: Do not mix with water
Dogs can make choline, but this is not sufficient
to meet all their bodies’ requirements. The rest
needs to be taken up from the diet in either free
or bound form, which reduces the need for methyl
groups supplied by methionine. Particularly dogs
receiving a high-fat diet require extensive lipid
transport by phosphatidylcholine. If an inadequate
amount of choline is made and consumed, liver
choline decreases rapidly [6] and the demand for
methyl groups cannot be met. In dogs, choline defi-
ciency results in liver dysfunction due to liver fat
accumulation [7].
In 2018, a survey of the Association for Pet Obesity
Prevention estimated that 60% of cats and 56% of
dogs in the United States were overweight or obese.
This has serious health consequences including fat
accumulation in the liver. A balanced, healthy diet
for dogs accompanied with regular physical activity
is essential for overall health. In addition, nutritional
supplementation strategies may help to reduce the
risk of weight-related disorders or optimize nutrition
for high-performance dogs. A mean to increase
choline intake in dogs is given in krill meal, which
provides choline in the bound form of phosphatidly-
choline [8, 9].
• Building of cell membranes
• Liver fat transport to prevent fatty liver
• Nerve impulse transmission in brain and muscles
• Providing methyl groups that maintain liver, heart and kidney health
CHOLINE
- 3 -
CHOLINE ISESSENTIAL FOR:
Because of the methyl group interrelationship, the
requirement for dietary choline supplementation
depends on the availability of methionine, betaine,
folate and vitamin B12. For example, betaine can
replace choline addition, since choline provides the
methyl group for betaine formation. Other factors
that determine the dietary choline requirement are
diet composition (protein, fat and carbohydrate
levels), as well as gender, age, activity level and
energy intake of dogs.
caused by special astaxanthin pigmentation spots
that can change in size and intensity [12]. The ability to
change pigmentation is important to balance darker
coloration for UV protection, when at the surface,
versus higher transparency for camouflage. The green
can be seen in their digestive system and results
from algae they eat. Krill can eat up to 20% of their
body weight per day, but they also have the ability to
shrink when starving and can survive without food
for up to 200 days.
Adult krill can get up to 6 cm long and has blue light
emitting spots on the body speculated to play a
role in mating, swarm formation or camouflage from
above when the sun shines on them. Surprisingly, the
krill genome is about 12 times the size of the human
genome. Antarctic krill is harvested in the wild and
then cooked, dried and ground. Whole krill gives a
powder of brownish-orange color rich in proteins,
choline, omega-3 fatty acids and astaxanthin.
Figure 3: Antarctic krill (Euphausia superba)
ANTARCTIC KRILL
- 4 -
Krill meal is made from Antarctic krill (Euphausia
superba), which is a crustacean related to shrimp (Fig.
3). But unlike shrimp that live on the ocean bottom,
krill aggregate in huge up to twenty-kilo-meter-long
swarms and move up and down the water column.
There are 85 different krill species around the world,
with Antarctic krill living in the Southern Ocean
around the frozen continent. Krill are transparent
with some red and green colorations. The red is
In humans, it was found that choline in the form of phospholipids is 12 times more efficient in raising blood choline
concentrations compared to choline chloride [10, 11].
While choline salt consumption in humans shows a 86% increase to a maximum plasma concentration after 30 minutes
returning to normal levels after four hours, phosphatidylcholine intake raises choline by 265% and takes 12 hours to decline [11].
Figure 5: Plasma choline levels
Figure 4: QRILL Pet ingredient
QRILL Pet is a functional
ingredient for pet food, rich
in phospholipid omega-3s,
choline, marine proteins and
astaxanthin. The choline found
in krill is all natural and in form
of phosphatidylcholine.
A previous study with krill meal was exploring it’s
benefits during an extreme endurance exercise, the
1600-km-long Iditarod dog sled race [13]. Iditarod
is perhaps the most extreme endurance exercise
performed by any domestic animal, and markers of
oxidative stress, inflammation and skeletal muscle
damage are known to be increased after the race [14].
The study reported that supplementation with krill
meal for 5 weeks before the race increased the
Omega-3 Index, which resulted in less inflammation
and muscle damage in the krill group in comparison
to a control group [13].
The latest study investigated if 8% dietary krill meal
inclusion for six weeks can increase the concen-
tration of choline and its metabolites in plasma of
Alaskan Huskies in comparison to a control group [15].
+52%
+18%
13,28
7,938,75
6,72
Baseline
Control QRILL Pet
End
KRILL MEAL RESEARCH
- 5 -
The results demonstrated that a nutritional strategy,
such as the addition of phosphatidylcholine to the
diet, can help to significantly increase plasma choline
concentrations (Fig. 5), which correlated with changed
concentrations of various choline metabolites (Fig. 6).
Chol
ine
(µm
ol/L
)
BETAINE (μmol/L) DIMETHYLGLYCINE (μmol/L)
BaselineBetaine promotes muscle function and plasma volume expansion.
Dimethylglycine is involved in glucose metabolism.BaselineEnd End
429,634,1
230,1 14,8
METHIONINE (μmol/L) HOMOCYSTEINE (μmol/L)
BaselineMethionine is an important methyl donor. Homocysteine is a risk factor for heart disease.
BaselineEnd
114,7
8,3106,2
10,1
End
Figure 6: Change in choline metabolites in plasma
These results could be of importance in a long-lasting
race setting, when a drop in plasma choline is expected
as seen in humans [16]. As a consequence of lowered
free plasma choline concentrations, the generation of
acetylcholine (a neurotransmitter important for
muscle function) might be reduced and thereby
negatively affect athletic performance. Noteworthy,
from a performance perspective, is also the significant
increase in betaine in the krill group, which is thought
to promote muscle function and plasma volume
expansion as shown after betaine supplementation in
humans [17]. The study also found a significant reduction
of plasma total homocysteine. Homocysteine is known
to increase the risk of endothelial cell injury and
cardiovascular disease in humans by increasing reactive
oxygen species and altering lipoprotein metabolism [18]. A similar correlation has been identified in dogs [19]. Another metabolic derivative of choline that was
significantly increased in the krill meal supplemented
dogs was dimethylglycine, which was suggested to have
possible protective effects on glucose metabolism [20].
The results of the krill meal supplementation study are encouraging to ensure that sufficient choline is available for optimal cell and body system functions. This is of particular interest for dogs that perform long-distance races or have health ailments affecting the liver, brain and heart.
Additionally, the omega-3 fatty acids from krill mayfurther benefit the health of dogs by decreasing inflammation. Low-grade chronic inflammation is a characteristic of various diseases such as obesity, cardiovascular disease, arthritis, diabetes, fatty liver disease and some cancers.
Similarly, the astaxanthin present in krill meal has anti-inflammatory and antioxidant properties through its ability to scavenge free radicals, thereby counteracting oxidative stress and damage to proteins, lipids and DNA. Since Antarctic krill is protected from overfishing by CCAMLR and Aker BioMarine Antarctic AS has obtained the Marine Stewardship Certification to prove sustainable fishing, QRILL Pet is an attractive feed ingredient that ensures optimal health of pets in a sustainable manner.
- 6 -
CONCLUSION
CHOLINE – AN ESSENTIAL NUTRIENT FOR DOGS
CHOLINE FORMS
HEALTH BENEFITS OF CHOLINE
SOURCES
HOW TO ENSURE YOUR DOG IS GETTING SUFFICIENT CHOLINE
Cholinechloride
Cholinebitartrate
Glycerophosphocholine
Citicoline
Phosphatidylcholine
REFERENCES
1. Burri L, Heggen K, Storsve AB (2019) Phosphatidylcholine from krill increases plasma choline and its metabolites in dogs, Veterinary World, 12(5): 671-676.
2. Wurtman, R.J., M.J. Hirsch, and J.H. Growdon, Lecithin consumption raises serum-free-choline levels. Lancet,1977. 2 (8028): p. 68-9.
increase of plasma choline
in dogs after 6 weeks on a
QRILL Pet diet
+52%
+18%
13,28
7,938,75
6,72
Baseline
Control QRILL Pet
End
52%
Choline from krill is a natural source of phosphatidylcholine (PC) . Another study has shown that consumption of choline in PC form also stays longer in the body. [2]
EGG
LIVER FISH
SOYBEAN
MEAT
WHEAT GERMKRILL
1700mgper kg dry matter
NRC recommendation
(2006) forall dogs
SOME OF THE MAJOR BENEFITS FOR HIGH PREFORMANCE DOGS
CARDIOVASCULARFUNCTION
COGNITIVEFUNCTION
OPTIMAL MUSCLEPERFORMANCE
DETOXIFICATION(METHYLATION)
IMPROVEDPERFORMANCE
LIVER
BRAIN & MEMORY
HEART
MUSCLE FUNCTION
REPRODUCTION
Recommended choline intake
WHY CHOLINE FROM KRILL?
The benefits of Phosphatidylcholine from krill in dog food
- 7 -
[1]
IMPROVEDHEALTH
ABOUT THE AUTHOR
Lena Burri, Ph.D., has been involved in funda- mental research and is together with co-authorscredited with several original protein discoveries. She has published scientific articles in leading journals, and contributed book chapters, review articles and peer-reviewed manuscripts on many subjects, including omega-3 fatty acids. Lena earned her Master of Science from the University of Basel (Switzerland) and her Ph.D. at the Ludwig Institute for Cancer Research (Switzerland).
Her post-doctoral education included stays at Melbourne University (Australia), University of British Columbia (Canada) and University of Bergen (Norway). She now works as R&D Directorfor Animal Nutrition and Health for Aker BioMarine.
THOMAS WÆRNER & CO.
Thomas Wærner is a professional dog musher and a member of the QRILL Pet Mushing Team, the world´s first professional long-distance dog sled team. His dog team participated in the krill meal supplementation study described in this whitepaper.
In 2019, Thomas and his dog team won both of the biggest long-distance sled dog races in Norway, Finnmark race and Femund race. During these races the teams cover distances up to 1200 km. Thomas also received the veterinarian award at the Finnmark race, which is handed out to themusher who demonstrated outstanding dog care and health.
He owns and runs the Berserk kennel in Synnfjell, Norway. His previous achievements include the Finnmark race in 2013, 3rd place at the Femund race in 2013 and “Rookie of the year” at the Iditarod race in 2015.
- 8 -
1. Food and Nutrition Board, I.o.M., Dietary Reference Intakes: Thiamin, Riboflavin, Niacin, Vitamin B-6, Vitamin B012, Pantothenic Acid, Biotin, and Choline. 1998, Washington, D.C.: National Academy of Sciences. p. 390-422.
2. Zeisel, S.H. and J.K. Blusztajn, Choline and human nutrition. Annu Rev Nutr, 1994. 14: p. 269-96.
3. Conlay, L.A., L.A. Sabounjian, and R.J. Wurtman, Exercise and neuromodulators: choline and acetylcholine in marathon runners. Int J Sports Med, 1992. 13 Suppl 1: p. S141-2.
4. Conlay, L.A., et al., Decreased plasma choline concentrations in marathon runners. N Engl J Med, 1986. 315(14): p. 892.
5. Marriott, B.M., Food Components to Enhance Performance: An evaluation of potential performance-enhancing food components for operational rations. 1994: National Academies Press.
6. Zeisel, S.H., T. Zola, and E. Pomfret, Effect of choline deficiency on S-adenosylmethionine and methionine concentrations in rat liver. Biochem J, 1989. 259(3): p. 725-729.
7. Best, C.H., J.M. Hershey, and M.E. Huntsman, The effect of lecithine on fat deposition in the liver of the normal rat. J Physiol, 1932. 75(1): p. 56-66.
8. Phleger, C.F., et al., Interannual and between species comparison of the lipids, fatty acids and sterols of Antarctic krill from the US AMLR Elephant Island survey area. Comp Biochem Physiol B Biochem Mol Biol, 2002. 131(4): p. 733-47.
9. Tou, J.C., J. Jaczynski, and Y.C. Chen, Krill for human consumption: nutritional value and potential health benefits. Nutr Rev, 2007. 65(2): p. 63-77.
10. Hirsch, M.J., J.H. Growdon, and R.J. Wurtman, Relations between dietary choline or lecithin intake, serum choline levels, and various metabolic indices. Metabolism, 1978. 27(8): p. 953-60.
11. Wurtman, R.J., M.J. Hirsch, and J.H. Growdon, Lecithin consumption raises serum-free-choline levels. Lancet, 1977. 2(8028): p. 68-9.
12. Auerswald, L., et al., Physiological and morphological colour change in Antarctic krill, Euphausia superba: a field study in the Lazarev Sea. J Exp Biol, 2008. 211(24): p. 3850-3858.
13. Burri, L., et al., Effects of dietary supplementation with krill meal on serum pro-inflammatory markers after the Iditarod sled dog race. Res Vet Sci, 2018. 121: p. 18-22.
14. Burr, J.R., et al., Serum biochemical values in sled dogs before and after competing in long-distance races. J Am Vet Med Assoc, 1997. 211(2): p. 175-9.
15. Burri, L., K. Heggen, and A. Storsve. Phosphatidylcholine from krill increases plasma choline and its metabolites in dogs, Vet World, 2019. 12(5): 671-676.
16. Jager, R., M. Purpura, and M. Kingsley, Phospholipids and sports performance. J Int Soc Sports Nutr, 2007. 4(1): p. 5.
17. Ismaeel, A., Effects of betaine supplementation on muscle strength and power: a systematic review. J Strength Cond Res, 2017. 31(8): p. 2338-2346.
18. Collaboration, H.S., Homocysteine and risk of ischemic heart disease and stroke: a meta-analysis. Jama, 2002. 288(16): p. 2015-2022.
19. Lee, C.-M., et al., Correlation between serum homocysteine concentration and severity of mitral valve disease in dogs. Am J Vet Res, 2017. 78(4): p. 440-446.
20. Magnusson, M., et al., Dimethylglycine deficiency and the development of diabetes. Diabetes, 2015. 64(8): p. 3010-3016.
- 9 -
REFERENCES
ABOUT QRILL PET
QRILL Pet is a product of Aker BioMarine, a leading biotechnology company developing and supplying krill- derived products for consumer health and animal nutrition. When developing our functional marine ingredient for pets, we kept our focus on the needs of pets and pet food formulators. Aimed at keeping pets healthy, QRILL Pet delivers long-chain omega-3 fatty acids, marine proteins, choline and the antioxidant astaxanthin. The unique distinction of krill omega-3s is that they are mainly bound to phospholipids.
QRILL Pet is currently the only MSC (Marine Stewardship Council) certified krill meal in the world. The ingredient comes form one of the most sustainable fisheries in the world and can be traced back to the exact catch location of the krill.
Aker BioMarine Antarctic AS | Qrillpet.com | Akerbiomarine.comQRILL Pet is a trademark of Aker BioMarine Antarctic AS, Norway © 2019 Aker BioMarine. All rights reserved.