4. Simkin AJ, Zhu C, Kuntz M, Sandmann G
(2003) Light-dark regulation of carotenoid
biosynthesis in pepper (Capsicum annuum)
leaves. J Plant Physiol 160:439–443
5. L€ atari K, Wu ¨st F, Hu ¨bner M et al (2015)
Tissue-specific Apocarotenoid glycosylation
contributes to carotenoid homeostasis in Arabidopsis leaves. Plant Physiol 168:1550–1562
6. Havaux M (2013) Carotenoid oxidation products as stress signals in plants. Plant J
79:597–606
7. Al-Babili S, Bouwmeester HJ (2015) Strigolactones, a novel carotenoid-derived plant hormone. Annu Rev Plant Biol 66:161–186
8. Ahrazem O, Rubio-Moraga A, Nebauer SG
et al (2015) Saffron: its Phytochemistry, developmental processes, and biotechnological prospects. J Agric Food Chem 63:8751–8764
9. Schaub P, Wuest F, Koschmieder J et al (2017)
Non-enzymatic β-carotene degradation in
(Provitamin A-biofortified) crop plants. J
Agric Food Chem. https://doi.org/10.1021/
acs.jafc.7b01693
10. Schaub P, Rodriguez-Franco M, Cazzonelli CI
et al (2018) Establishment of an Arabidopsis
callus system to study the interrelations of biosynthesis, degradation and accumulation of
carotenoids. PLoS One 13:e0192158
11. Ramel F, Mialoundama AS, Havaux M (2013)
Nonenzymic carotenoid oxidation and photooxidative stress signalling in plants. J Exp Bot
64:799–805
12. Walter MH, Strack D (2011) Carotenoids and
their cleavage products: biosynthesis and functions. Nat Prod Rep 28:663–692
13. Beisel KG, Jahnke S, Hofmann D et al (2010)
Continuous turnover of carotenes and chlorophyll a in mature leaves of Arabidopsis thaliana
revealed by 14CO2 pulse-chase labeling. Plant
Physiol 152:2188–2199
14. Beisel KG, Schurr U, Matsubara S (2011)
Altered turnover of β-carotene and Chl a in
Arabidopsis leaves treated with lincomycin or
norflurazon. Plant Cell Physiol 52:1193–1203
15. Breitenbach J, Zhu C, Sandmann G (2001)
Bleaching herbicide norflurazon inhibits
phytoene desaturase by competition with
the cofactors. J Agric Food Chem 49:
5270–5272
16. Koschmieder J, Fehling-Kaschek M, Schaub P
et al (2017) Plant-type phytoene desaturase:
functional evaluation of structural implications.
PLoS One 12:e0187628
17. Brausemann A, Gemmecker S, Koschmieder J
et al (2017) Structure of Phytoene Desaturase
provides insights into herbicide binding and
reaction mechanisms involved in carotene desaturation. Structure 25(8):1222–1232.e3
18. Welsch R, Zhou X, Yuan H et al (2017) Clp
protease and OR directly control the proteostasis of phytoene synthase, the crucial enzyme
for carotenoid biosynthesis in Arabidopsis. Mol
Plant 11:149–162
19. Zhou X, Welsch R, Yang Y et al (2015) Arabidopsis OR proteins are the major posttranscriptional regulators of phytoene synthase in
controlling carotenoid biosynthesis. Proc Natl
Acad Sci 112:3558–3563
20. Rodrı ´guez-Villalo ´ n A, Gas E, Rodrı ´guez-Concepcio ´ n M (2009) Phytoene synthase activity
controls the biosynthesis of carotenoids and
the supply of their metabolic precursors in
dark-grown Arabidopsis seedlings. Plant J
60:424–435
21. Rodrı ´guez-Villalo ´ n A, Gas E, Rodrı ´guezConcepcio ´ n M, Rodriguez-Villalon A (2009)
Colors in the dark: a model for the regulation
of carotenoid biosynthesis in etioplasts. Plant
Signal Behav 4:965–967
22. A ´ lvarez D, Voß B, Maass D et al (2016) Carotenogenesis is regulated by 5’UTR-mediated
translation of Phytoene synthase splice variants.
Plant Physiol 172:2314–2326
23. Britton G, Liaaen-Jensen S, Pfander H (1994)
Carotenoids—Volume 1 B: Spectroscopy, 1st
edn. Birkh€ auser, Basel
24. Britton G, Liaaen-Jensen S, Pfander H (1994)
Carotenoids—volume 1 a: isolation and analysis, 1st edn. Birkh€ auser, Basel
25. Hoa TTC, Al-Babili S, Schaub P et al (2003)
Golden Indica and japonica rice lines amenable
to deregulation. Plant Physiol 133:161–169
Quantification of Carotenoid Pathway Flux
291
(2003) Light-dark regulation of carotenoid
biosynthesis in pepper (Capsicum annuum)
leaves. J Plant Physiol 160:439–443
5. L€ atari K, Wu ¨st F, Hu ¨bner M et al (2015)
Tissue-specific Apocarotenoid glycosylation
contributes to carotenoid homeostasis in Arabidopsis leaves. Plant Physiol 168:1550–1562
6. Havaux M (2013) Carotenoid oxidation products as stress signals in plants. Plant J
79:597–606
7. Al-Babili S, Bouwmeester HJ (2015) Strigolactones, a novel carotenoid-derived plant hormone. Annu Rev Plant Biol 66:161–186
8. Ahrazem O, Rubio-Moraga A, Nebauer SG
et al (2015) Saffron: its Phytochemistry, developmental processes, and biotechnological prospects. J Agric Food Chem 63:8751–8764
9. Schaub P, Wuest F, Koschmieder J et al (2017)
Non-enzymatic β-carotene degradation in
(Provitamin A-biofortified) crop plants. J
Agric Food Chem. https://doi.org/10.1021/
acs.jafc.7b01693
10. Schaub P, Rodriguez-Franco M, Cazzonelli CI
et al (2018) Establishment of an Arabidopsis
callus system to study the interrelations of biosynthesis, degradation and accumulation of
carotenoids. PLoS One 13:e0192158
11. Ramel F, Mialoundama AS, Havaux M (2013)
Nonenzymic carotenoid oxidation and photooxidative stress signalling in plants. J Exp Bot
64:799–805
12. Walter MH, Strack D (2011) Carotenoids and
their cleavage products: biosynthesis and functions. Nat Prod Rep 28:663–692
13. Beisel KG, Jahnke S, Hofmann D et al (2010)
Continuous turnover of carotenes and chlorophyll a in mature leaves of Arabidopsis thaliana
revealed by 14CO2 pulse-chase labeling. Plant
Physiol 152:2188–2199
14. Beisel KG, Schurr U, Matsubara S (2011)
Altered turnover of β-carotene and Chl a in
Arabidopsis leaves treated with lincomycin or
norflurazon. Plant Cell Physiol 52:1193–1203
15. Breitenbach J, Zhu C, Sandmann G (2001)
Bleaching herbicide norflurazon inhibits
phytoene desaturase by competition with
the cofactors. J Agric Food Chem 49:
5270–5272
16. Koschmieder J, Fehling-Kaschek M, Schaub P
et al (2017) Plant-type phytoene desaturase:
functional evaluation of structural implications.
PLoS One 12:e0187628
17. Brausemann A, Gemmecker S, Koschmieder J
et al (2017) Structure of Phytoene Desaturase
provides insights into herbicide binding and
reaction mechanisms involved in carotene desaturation. Structure 25(8):1222–1232.e3
18. Welsch R, Zhou X, Yuan H et al (2017) Clp
protease and OR directly control the proteostasis of phytoene synthase, the crucial enzyme
for carotenoid biosynthesis in Arabidopsis. Mol
Plant 11:149–162
19. Zhou X, Welsch R, Yang Y et al (2015) Arabidopsis OR proteins are the major posttranscriptional regulators of phytoene synthase in
controlling carotenoid biosynthesis. Proc Natl
Acad Sci 112:3558–3563
20. Rodrı ´guez-Villalo ´ n A, Gas E, Rodrı ´guez-Concepcio ´ n M (2009) Phytoene synthase activity
controls the biosynthesis of carotenoids and
the supply of their metabolic precursors in
dark-grown Arabidopsis seedlings. Plant J
60:424–435
21. Rodrı ´guez-Villalo ´ n A, Gas E, Rodrı ´guezConcepcio ´ n M, Rodriguez-Villalon A (2009)
Colors in the dark: a model for the regulation
of carotenoid biosynthesis in etioplasts. Plant
Signal Behav 4:965–967
22. A ´ lvarez D, Voß B, Maass D et al (2016) Carotenogenesis is regulated by 5’UTR-mediated
translation of Phytoene synthase splice variants.
Plant Physiol 172:2314–2326
23. Britton G, Liaaen-Jensen S, Pfander H (1994)
Carotenoids—Volume 1 B: Spectroscopy, 1st
edn. Birkh€ auser, Basel
24. Britton G, Liaaen-Jensen S, Pfander H (1994)
Carotenoids—volume 1 a: isolation and analysis, 1st edn. Birkh€ auser, Basel
25. Hoa TTC, Al-Babili S, Schaub P et al (2003)
Golden Indica and japonica rice lines amenable
to deregulation. Plant Physiol 133:161–169
Quantification of Carotenoid Pathway Flux
291