Annals of Applied Biology ISSN 0003-4746
RESEARCH ARTICLE
Post-flowering photoperiod has marked effects on fruit
chemical composition in red raspberry (Rubus idaeus)
S.P. Mazur
1,2
, A. Sønsteby
1
, A.-B. Wold
2
, A. Foito
3
, S. Freitag
3
, S. Verrall
4
, S. Conner
3
, D. Stewart
1,3
&
O.M. Heide
5
1 Bioforsk, Norwegian Institute for Agricultural and Environmental Research, Kapp, Norway
2 Department of Plant and Environmental Sciences, Norwegian University of Life Sciences, Ås, Norway
3 Environment and Biochemical Sciences, James Hutton Institute, Dundee, UK
4 Information and Computational Sciences, James Hutton Institute, Dundee, UK
5 Department of Ecology and Natural Resource Management, Norwegian University of Life Sciences, Ås, Norway
Keywords
Anthocyanins; antioxidant capacity; ellagic acid;
fruit quality; harvest time; metabolite profile;
organic acids; photoperiod; polyphenols; Rubus
idaeus; sugars; vitamin C.
Correspondence
S.P. Mazur, Bioforsk, Norwegian Institute for
Agricultural and Environmental Research,
NO-2849 Kapp, Norway.
Email: sebastian.mazur@bioforsk.no
Received: 18 December 2013; revised version
accepted: 14 July 2014.
doi:10.1111/aab.12153
Abstract
The effect of photoperiod on nutritional quality and chemical composition of
‘Glen Ample’ red raspberry has been studied under controlled environment
conditions. Long day (LD) conditions during fruit growth and maturation signif-
icantly increased fruit concentrations of ascorbic acid, total phenolics, organic
acids such as malic, quinic and ellagic acids, as well as the ferric reducing activ-
ity power, while reducing the concentrations of soluble solids, sucrose and the
sugar/acid ratio. The levels of total anthocyanin and oxalic acid were not signif-
icantly affected by photoperiod whereas semi-quantitative analysis indicated a
significant increase in the levels of some ellagitannins. The same responses were
found whether LD was established by day-length extension with low-intensity
light or by night interruption, thus demonstrating that the observed changes in
nutritional quality of raspberry fruit was mediated through specific photoperi-
odic mechanism(s) independently of the daily light energy supply.
Introduction
Fruits, and berries in particular, are rich dietary sources of
a range of vitamins, minerals and other phytochemicals
of interest for human nutrition and health (Sun et al.,
2002, and ref. therein; Rao & Snyder, 2010). Traditionally,
L-ascorbic acid (L-AA, vitamin C) has been of predomi-
nant interest because of its antiscorbutic properties, but
more recently dietary phenolic compounds have received
increasing interest due to their potential dietary health
benefits (Ness & Powles, 1997; Mullen et al., 2002; Reddy
& Katan, 2004; Stanner et al., 2004; López et al., 2010)
which were thought to be linked to their direct antiox-
idant capacities (AOC). However, due to the reported
limited bioavailabilities of such phenolic antioxidants
(Manach et al., 2005; Williamson & Manach, 2005), this
is now less clear with mechanisms such as induction
of endogenous antioxidant systems, cyclooxygenase
inhibition and upregulation of endothelial nitric oxide
synthase (eNOS) proposed (Smoliga et al., 2011). Public
awareness of the potential health benefits of increased
intake of fruit and vegetables has increased consumer
demands for fresh fruit and associated product, and has
intensified research activity on the effects of genetic and
environmental factors on fruit chemical composition
(Atkinson et al., 2005; and references therein).
In the Nordic countries, there has been a long-held
notion that the concentration of L-AA and other
health-related phytochemicals in plants are increas-
ing with the latitude of production (Jaakola & Hohtola,
2010; Remberg et al., 2010), and there is also some sci-
entific evidence from both fruit and vegetable research
to support this (Hårdh & Hårdh, 1977; Hårdh et al., 1977;
Davik et al., 2006; Jaakola & Hohtola, 2010; Remberg
et al., 2010). It is believed that the long days (LDs), high
total radiation and cool temperatures of the high-latitude
summer may increase the synthesis of aromatic com-
pounds and vitamins such as L-AA (Jaakola & Hohtola,
2010). Since all these environmental factors, as well
as light quality, change with increasing latitude, it is
difficult to determine which factor(s) are the causal
Ann Appl Biol (2014) 1
© 2014 Association of Applied Biologists