Acta Sci. Pol., Hortorum Cultus 13(5) 2014, 91-105
DIFFERENCES IN THE STRUCTURE OF FRUIT BUDS
IN TWO APPLE CULTIVARS WITH PARTICULAR
EMPHASIS ON FEATURES RESPONSIBLE FOR FRUIT
STORABILITY AND QUALITY
University of Life Sciences in Lublin
Abstract. ‘Jonagold’ and ‘Szampion’ are winter apple cultivars, whose fruits are suitable
for long-term storage. However, fruits of these cultivars differ markedly in the type of the
surface and the rate and volume of water transpiration, which is manifested in fruit quality
after storage and the length of apple shelf life. A majority of factors responsible for fruit
quality and storability are genetically conditioned traits that are mainly developed before
fruits reach harvest maturity or still develop during the storage period. The micromor-
phology, anatomy, and ultrastructure of 21-day-old fruit buds of the ‘Jonagold’ and
‘Szampion’ were examined using light microscopy as well as scanning and transmission
electron microscopy. The analyses were particularly focused on the traits that determine
fruit firmness and storability, which contribute to long-term storage capacity. It was found
that the fruit buds in both cultivars differed significantly in the number of trichome scars
and stomata on the fruit surface, the thickness of the hypodermis layer and the hypoder-
mis cell walls, and in the content of phenolic compound deposits. At the fruit bud stage,
the following features related to increased or decreased fruit firmness and storability were
observed: platelet crystalline wax, cuticle microcracks, stomata and trichome scars, and
presence of phenolic compounds.
cuticle and epicuticular wax, microcracks, phenol compounds
Apple cultivars are hybrids of Malus domestica (Borkh). They differ in many traits,
e.g. the timing of reaching harvest and consumption maturity and long-term shelf life of
fruits, which are largely dependent on the genetic background as well as on climatic and
storage conditions [Rejman 1994, Kruczy ska 2008]. Apple fruits develop from the
ovary of the flower and from the floral tube (receptacle). The development proceeds in
Corresponding author: Agata Konarska, Department of Botany, University of Life Sciences in
Lublin, Akademicka 15, 20-950 Lublin, Poland, e-mail: firstname.lastname@example.org
two distinct stages, and the fruit growth curve has a form of a sigmoid curve [Miller et
al. 1987, Westwood 1995]. During the development, the fruit size and weight increase
systematically, while firmness and acidity decline [Atay et al. 2010]. The first stage of
fruit development, i.e. the so-called fruit bud stage, is characterised by very intensive
cell divisions (up to 8–10 weeks after anthesis). During this period, fruits are firm and
contain a lot of starch, tannins and acids; therefore, they are tart and sour. In the second
stage, between mid-June and harvest time, the fruit volume increases through enlarge-
ment of the intercellular spaces and cell volume. Additionally, accumulation of food
reserves occurs, large vacuoles emerge in the cells, and the water content in fruits in-
creases in this stage. Physical changes in fruits are accompanied by chemical processes
leading to reduction of the content of tannins and intensive accumulation of pigments,
sugars, organic acids, fatty substances and pectins, volatile substances, vitamins, and
minerals [Pieni ek 2000, Atay et al. 2010].
Fruits of different varieties of Malus vary with the the type of the fruit surface and
peel structure and exhibit variation in the intensity of transpiration [Belding et al. 1998,
Veraverbake et al. 2001a]. These traits are primarily associated with the amounts of
cuticular (intra- and epicuticular) waxes as well as the number and depth of epidermal
microcracks [Gordon et al. 1998, Maguire et al. 2000, Veraverbake et al. 2003b, Konar-
ska 2012]. A great impact on fruit transpiration is also exerted by the number of
“opened” (active) stomata and lenticels per unit surface area and, to a small extent, by
cuticle thickness and the number of hypodermis layers [Veraverbake 2001b, 2003b,
Homutová and Blažek 2006]. These features prevent or promote fruit water loss, which
is reflected in their firmness, post-storage quality (attractiveness), and the length of shelf
life [Riederer and Schreiber 1995, Czernyszewicz 2007]. The peel morphology play
also an important role in determining the distributions of water, carbohydrates, and
nutrients inside the fruit [Cieslak et al. 2013]. According to Khanal and Knoche 
the epidermal and hypodermal cell layers represent the structural backbone of an apple
peel during pre- and postharvest development, whereas cutical membrane microcrack-
ing has limited relevance to the overall mechanical properties of the peel. Recent re-
search shows that, similar to the cuticle layer, the outer layer of the cuticle (cuticle
proper) contains polysaccharides (cellulose and pectin), which can affect the rheological
properties of cuticles and may actively contribute to the bi-directional transport of water
and solutes [Guzmán et al. 2014]. Moreover, optical coherence tomography which is
a new non-destructive technique to visualize subsurface structures of materials, can be
used to demonstrate peel structural differences between apples, as well as to measure
structural changes that occur during storage [Verboven et al. 2013].
In previous studies, the author of the paper found that, the peels of two apple culti-
vars ‘Jonagold’ and ‘Szampion’ differed in many quantitative and qualitative traits of
their micromorphology, anatomy, and ultrastructure at the harvest and consumption
maturity stage [Konarska 2013]. The most substantial differences between the cultivars
involved the quantity and forms of epicuticular wax and the total wax weight; the depth
of cuticular microcracks; the number of stomata and lenticels; cuticle thickness and the
size of epidermal cells; the thickness of the hypodermis layer; and the presence of phe-
nolic compounds. The extensive literature concerning apple development and structure
does not provide detailed information about the morphology, anatomy, and ultrastruc-
ture of apple fruit buds or the mechanism and time of development of traits responsible
major qualitative fruit traits may be important for growers, who could control and mod-
ify the trait development through application of appropriate fertilisation and/or irrigation
and other agricultural treatments. Therefore, the aim of the present study was to analyse
the structure of 21-day-old ‘Jonagold’ and ‘Szampion’ fruit buds at the intensive cell
division stage and to present differences in the structure of the covering layer in these
cultivars at the micromorphological, tissue, and cellular level. Particular attention was
paid to the occurrence of traits that have an impact on subsequent quality and firmness
MATERIAL AND METHODS
21-day-old (21 days after anthesis) fruit buds (ca. 1-cm diameter) of ‘Jonagold’ and
‘Szampion’ apples were collected on May 15–20, 2012 in a commercial, conventionally
managed orchard near Lublin. Trees of the analysed cultivars grew in close proximity
and identical climate and soil conditions. 20 fruit buds were sampled from the crowns of
5 randomly chosen trees of both cultivars. Further analyses were carried out on peel-
comprising fragments of fruits sampled from their equatorial part.
Scanning electron microscopy (SEM). The fruit buds were carefully transported
to the laboratory to avoid damage or destruction of their surface wax layer. Next,
4 × 4 × 1-mm peel fragments were sampled from 5 fruits of each cultivar. Since fixation
of the material for SEM induces changes in the structure and destruction of the epicu-
ticular wax layer [Reed 1982], freshly sampled material was carefully mounted onto
stubs, sputter-coated with gold, and examined “live” under a TESCAN/VEGA LMU
(Tescan, USA) scanning electron microscopy at an accelerating voltage of 30 kV. Quan-
tity of wax platelets was roughly determined, while the the length and the number of
stomata and trichome scars within an area of 1 mm
of the epidermis were assessed
to the fruit axis were made through fresh peel of 10 fruit buds of ‘Jonagold’ and ‘Szam-
pion’. Further, the samples were stained with Sudan III (a saturated ethanol solution of
Sudan III) to visualize lipophilic substances in cuticle, with Lugol’s iodine in order to
detect starch, and with FeCl
to detect phenolic substances. Later, the samples were
light microscope where the thickness of the cuticle (at the mid-width of a randomly
chosen epidermal cell), the height of the epidermal cells, the number of layers of hypo-
dermis and its overall thickness, the thickness of hypodermis cell walls, and the thick-
ness of 3 layers of the parenchyma located under the hypodermis were determined in
five places under a Nikon SE 102 light microscopy. Hand-cut samples obtained from
fresh material were also observed with a stereoscopic Nikon Eclipse 90i microscope
combined with an UV filter set comprising the wavelength of EX 330–380 nm stimulat-
ing autofluorescence of cuticle and chlorophyll in order to analyse the distribution
of that substances. Images were obtained by using a digital camera (Nikon Fi1) and
NIS-Elements Br 2 software, respectively a Zeiss Axiolmager Z1 fluorescence micro-
scope equipped with an AxioCam MR digital camera.
0.7-μm semi-thin sections were also prepared from fragments of the fruit buds,
which were stained with 1% methylene blue with 1% azur II in a 1% aqueous solution
of sodium tetraborate. The material was fixed and embedded in synthetic resin with the
standard method used in transmission electron microscope (see below). Sections were
observed by means of a Nicon Eclipse 90i microscopy.
5 fruit buds of ‘Jonagold’ and ‘Szampion’ were fixed in 2% paraformaldehyde and
2.5% glutaraldehyde buffered at pH 7.4 in 0.1 M cacodylate buffer. Fixation was per-
formed at room temperature for two hours, followed by 12 hr at 4ºC. When fixed, the
samples were rinsed with 0.1 M cacodylate buffer at 4ºC for 24 hr and then treated with
. After passage through increasing concentrations of propylene oxide in etha-
nol and finally through pure propylene oxide, the samples were embedded for 12 hr in
Spurr Low Viscosity resin at 70ºC [Spurr 1969]. Subsequently, ultrathin sections
(70 nm thick) obtained using the Reichert Ultracut-S ultramicrotome (Vienna, Austria)
and a glass knife were transferred to re-distilled water and stained with a 0.5 M aqueous
solution of uranyl acetate and lead citrate [Reynolds 1963]. Images were observed and
recorded using the FEI Technai G2 Spirit Bio TWIN transmission electron microscopy
at an accelerating voltage of 120 kV. Images were captured using a Megaview G2
Olympus Soft Imaging Solutions camera.
Data were analysed by one-way analysis of variance (ANOVA) and Tukey’s multiple
range test for comparison of means, using software Statistica 7. The difference was
considered statistically significant at the level of P < 0.05.
ken off, unicellular, ca. 1-mm-long non-glandular trichomes (figs 1A, C). The epidermis
among the trichomes exhibited few trichome scars, i.e. traces left by broken off non-
glandular trichomes (figs 1D–E), and stomata in various developmental stages showing
different degrees of stomatal opening (figs 1F, G). In both cultivars, the sizes of stomata
and trichome scars were comparable, but their number per unit surface area was by ca.
30% greater in ‘Jonagold’ (tab. 1). The cuticle of both cultivars exhibited microcracks
of varied depths: superficial, which were more abundant, and deeper microcracks,
which occurred sporadically and resembled a fastened zipper (fig. 1H). Moreover, the
cuticle surface displayed vertically and horizontally oriented crystalline wax platelets
(figs 2A–D). In ‘Jonagold’, the wax platelets were more ordered and more numerous
than in the ‘Szampion’ cultivar. Additionally, a majority of the platelets were vertically
or obliquely arranged (fig. 2A, B).
LM. The surface layer of the 21-day-old ‘Jonagold’ and ‘Szampion’ fruit buds was
composed of a single-layered epidermis covered by a cuticle layer and several hypo-
dermis layers (figs 3A–F).
Fig. 1. Epidermis surface of the fruit buds in the ‘Jonagold’ (A, C, D, F, G) and ‘Szampion’
cultivars (B, E, H); A, C – fragments of the epidermis surface with numerous subulate
non-glandular trichomes; B – a fruit bud of ‘Szampion’ in stereoscopic microscopy.
C – visible trichomes, stomata (arrows) and trichome scars (arrowheads); D, E – trichome
scars (arrows) visible on the fruit bud surface; D – note a breaking off non-glandular
trichome; F, G –stomata in different stages of development; H – microcrack (arrows) and
trichome scar (arrowhead)
Fig. 2. SEM. Fragments of the epidermis surface of the fruit buds ‘Jonagold’ (A, B) and ‘Szam-
rangement and a greater number of vertical and inclined wax plates in ‘Jonagold’
Epidermis covering the fruit buds in both cultivars had a nature of a meristematic
nal elongated cells, particularly in the ‘Szampion’ cultivar (figs 3C, D). The height of
epidermal cells was by 8% greater in ‘Jonagold’ (tab. 1). The epidermis was covered by
a different-thickness cuticle layer which was stained orange-red by Sudan 3 (not shown)
and exhibiting light blue fluorescence under UV light (fig. 3E). The mean cuticle thick-
ness was by 11% greater in the ‘Szampion’ cultivar (tab. 1). The epidermis of both
cultivars had stomata with large air chambers underneath (fig. 3F). In the ‘Jonagold’
cultivar, the hypodermis layer was by 11% thicker than in ‘Szampion’ (tab. 1) and con-
sisted of 5–6 layers of the rectangular outline collenchyma cells with distinctly thick-
Fig. 3. LM. Fragments of the cross-sections through the ‘Jonagold’ (A, B, E, F) and ‘Szam-
pion’(C–D) surface layer of the fruit bud. A, B – in hypodermis and parenchymatic cells
visible large deposits of phenolic substances (arrowheads) and thickened tangential cell
walls in the hypodermis cells (arrows with two heads). Note epidermis cells after mitotic
divisions (asterisks); B – visible chloroplasts (arrows) in the hypodermis; C, D – visible
chloroplasts (arrows) (D) and cells after mitotic divisions in the epidermis, hypodermis
and parenchyma layer (asterisks); E – visible blue-fluorescent cuticle and red – fluores-
cent chloroplasts (fluorescence microscopy); F – visible stoma (S) with the air space (as-
terisk). Note chloroplasts (arrows) and deposits of phenolic substances (arrowheads) in
the hypodermis cells; C – cuticle, E – epidermis, H – hypodermis, P – parenchyma
bud stage; A, B – visible cuticle composed of lamellar cuticle proper (CP) and a reticulate
cuticular layer (CL); A, C – plastids (P) with starch grains visible in the epidermis and
hypodermis cells; D – chloroplasts (Ch) with starch grains and electron-dense deposits of
phenolic substances (arrowheads) visible in the hypodermis cells; E – deposit of phenolic
substances visible in the vacuole; N – nucleus, M – mitochondrion, V – vacuoles,
CW – cell wall
ened tangential walls (tab. 1, figs 3A, B). In turn, the hypodermis cells in the ‘Szam-
the differentiation stage and had an oval outline characteristic of parenchymatic cells
(tab. 1, figs 3C, D). The diameters of parenchymal cells located below the hypodermis
were comparable in both cultivars (tab. 1, figs 3A, C). Likewise in the epidermis, cells
after mitotic divisions were also observed in hypodermal and parenchymal layers, par-
ticularly in the ‘Szampion’ cultivar (figs 3C, D). Numerous chloroplasts that produced
red fluorescence under the fluorescence microscopy (fig. 3E) and contained starch (re-
action with IKI) were visible in the cytoplasm of these tissues in both cultivars (figs 3B,
D, F). Additionally, the hypodermis and parenchymal cells in the ‘Jonagold’ fruit buds
contained oval deposits of phenolic compounds (figs 3A, B, F) characterised by dark
brown staining in FeCl
Parameters (n = 10)
Number of stomata and trichome scars (per mm
18 ±4.0 b
Length of stomata pores (μm)
27.8 ±3.3 a
30.1 ±2.9 a
Length of trichome scars (μm)
23.32 ±4.6 a
22.56 ±4.2 a
Thickness of cuticle (μm)
8.64 ±0.7 a
9.77 ±0.9 a
Height of the epidermis cells (μm)
22.93 ±1.7 a
21.1 ±1.2 a
Number of the hypodermis layers
6 ±1.0 a
4 ±1.0 a
Thickness of the hypodermis layer (μm)
85.51 ±9.7 a
76.18 ±5.8 b
Thickness of the hypodermis cell walls
3.65 ±1.0 a
0.09 ±0.4 b
Thickness of the three parenchyma layers (μm)
65.71 ±11.1 a
67.98 ±6.0 a
Total thickness of peel (μm)
114.0 ±23.6 a
106.3 ±18.4 a
Values are mean ±SD (standard deviation). The same letters within a row mean no statistically
parenchymal cells were found between the analysed cultivars. The cuticle on the surface
of the fruit buds was composed of two layers: a substantially larger internal reticulate
layer, the so-called cuticular layer and an external lamellate layer, the so-called cuticle
proper, accounting for ca. 8–10% of the total cuticle thickness (figs 4A, B). The epi-
dermal cells exhibited a thin layer of parietal cytoplasm with visible mitochondria, and
plastids containing starch grains (figs 4A, C). Similarly, the cytoplasm of hypodermal
and parenchymal cells had plastids containing starch grains, whereas the vacuoles in the
‘Jonagold’ cultivar exhibited numerous, large electron-dense deposits of phenolic com-
pounds (figs 4D, E). They usually adhered to the tonoplast and had different sizes.
Most traits related to fruit quality and firmness are genetically conditioned [Faust
and Shear 1972a]. In the fruits of ‘Jonagold’ and ‘Szampion’, these traits were fully
developed at harvest and consumption maturity [Konarska 2013]. 3-week-old fruit buds
of the ‘Jonagold’ and ‘Szampion’ cultivars exhibited most features associated with the
protective function of the surface covering layer. Differences in the structure of the fruit
buds between the cultivars were visible primarily at the level of anatomy and micro-
morphology and were less evident than in the stage of harvest and consumption matur-
Babos et al.  and Zamorsky  report that protection of the fruit interior is
hypodermis. In the early stage of ‘Jonagold’ and ‘Szampion’ fruit development, the
protective role of this layer is additionally strengthened by abundant non-glandular
trichomes densely distributed in the fruit bud epidermis. Similar non-glandular
trichomes with a similar function were found to cover fruits of other plant species in
different developmental stages [Bain 1961, Miller 1984, Bednorz and Wojciechowicz
2009, Celano et al. 2009]. In 21 day-old ‘Jonagold’ and ‘Szampion’ fruit buds only few
trichome scars remaining after broken off trichomes were visible. Maguire et al. 
and Veraverbake et al. [2003b] report that trichome scars, stomata and lenticels, present
in the fruit epidermis at all developmental stages facilitate gas exchange and promote
fruit transpiration, thereby contributing to wilting, softening and quality deterioration
during storage and shelf life. However, the total number of trichome scars and stomata
in the ‘Szampion’ fruit buds was lower than that in ‘Jonagold’. As reported by Konarska
, this relationship persisted in the consumption maturity stage, although ‘Szam-
pion’ fruits exhibit worse and shorter storability. According to Veraverbake et al.
[2003a], at the consumption maturity stage, approximately 60% of ‘Jonagold’ lenticels
are “closed” (non-transpiring), while only “opened” lenticels promote transpiration.
A vast majority of the ‘Szampion’ lenticels may have been “opened”; yet, no such in-
vestigations have been conducted. The author of the present study considers that the
intensity of fruit transpiration is dependent on a set of several water-loss promoting
traits rather than on a single feature.
The surface of the fruit buds in both cultivars examined showed few microcracks
present mostly on the surface cuticle layers. The presence of the microcracks indicated
the onset of the cell expansion process, although cells after mitotic divisions were still
found by the author in all the layers of the fruit bud tissues, i.e. the epidermis, hypoder-
mis, and parenchyma. Microcracks appearing in the cuticle may indicate more rapid
expansion of the volume and turgor of epidermal and internal fruit cells, while the
amount of cuticle per fruit is constant [Faust and Shear 1972a, Roy et al. 1994, Knoche
et al. 2004]. Similar results were obtained by Harker and Ferguson , who found
that the fruit bud stage in apples is a period of dynamic cell divisions, particularly in the
epidermis, and the beginning of intensive cell growth. According to many authors, mi-
crocracks enhance water loss and reduce fruit firmness and weight [H hn 1990, Lau and
Lane 1998, Maguire et al. 1999, De Bellie 2000, Link et al. 2004].
The surface of the fruit buds in the cultivars examined was covered by a layer of
more abundant and ordered in the ‘Jonagold’ cultivar. Unlike the trichome scars, sto-
mata as well as microcracks, cuticular waxes are an efficient protective barrier against
excessive transpiration [Faust and Shear 1972b, Babos et al. 1984, Roy et al. 1994,
Belding et al. 1998, Veraverbeke et al. 2001a, b]. The number of wax platelets increases
together with fruit maturation, reaching a maximum value during the storage period,
particularly in varieties with a greasy and smooth peel, which was observed by Konar-
ska  in ‘Jonagold’ fruits stored in a controlled-atmosphere storehouse for
6 months. Koch et al.  and Curry  suggest that production of the highest
possible numbers of vertically oriented platelets is especially important for reduction of
transpiration, since this form of wax promotes healing and “repair” of microcracks.
According to Curry [2001, 2005] and Müller , apple trees employ a ‘Tear and
Repair’ mechanism involving continuous synthesis of epicuticular waxes and closure of
microcracks appearing along with fruit growth.
The surface of the fruit buds of the cultivars examined had a characteristic reticulate-
lamellate cuticle, whose layer was thicker in the ‘Szampion’ cultivar. As shown by
Konarska , ‘Szampion’ fruits were characterised by greater weight loss and more
intensive transpiration during the storage period. A thicker cuticle does not restrict the
decline in fruit firmness and their better quality after storage. Similar results concerning
the role of the cuticle in fruits of other apple varieties were obtained by Riederer and
Schreiber  and Knoche et al. .
Numerous deposits of phenolic compounds were found in the hypodermis and par-
enchymal cells in the ‘Jonagold’ fruit buds; these, however, were not observed in
‘Szampion’. Similarly, the deposits were visible, although in smaller numbers, only in
the harvest and consumption maturity stage in ‘Jonagold’ [Konarska 2013]. The results
obtained by the author correspond to the findings of Mehrabani and Hassanpouraghdam
, who reported that the content of phenolic compounds was higher in younger
fruits than in the harvest maturity stage. Additionally, a varied content of polyphenols in
different apple and pear cultivars has been described by other researchers [Solovchenko
and Schmitz-Elberger 2003, Drogoudi et al. 2008, ata et al. 2009]. Absence of poly-
phenols in the ‘Szampion’ cultivar may be one of the causes of the poorer quality and
storability of its fruits, as evidenced by literature data indicating that presence of poly-
phenols improves and extends fruit storage by increasing resistance to pathogens [Garry
et al. 1995, Lattanzio et al. 2001, Cheynier 2005].
1. The fruit buds in the ‘Jonagold’ and ‘Szampion’ cultivars differed significantly in
the number of trichome scars and stomata, thickness of hypodermis layer and hypoder-
mis cell walls, as well as the quantities of deposits of phenolic compounds.
2. The following factors exert an impact on the fruit quality and storability in the
fruit bud stage: trichome scars and stomata, microcracks, crystalline wax platelets and
This work was supported by the Ministry of Science and Higher Education of Po-
land as part of the statutory activities of the Department of Botany, University of Life
Sciences in Lublin.
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RÓ NICE W STRUKTURZE ZAWI ZKÓW OWOCÓW DWÓCH
UPRAWNYCH ODMIAN JAB ONI ZE SZCZEGÓLNYM
UWZGL DNIENIEM CECH DECYDUJ CYCH O ICH TRWA O CI
I JAKO CI
Streszczenie. ‘Jonagold’ i ‘Szampion’ nale do zimowych odmian jab oni, których owo-
ce s przystosowane do d ugotrwa ego przechowywania. Jednak owoce ró ni si wyra -
nie rodzajem powierzchni oraz tempem i ilo ci transpirowanej wody, co przek ada si na
jako owoców po wyj ciu z przechowalni oraz na d ugo ycia jab ek na pó ce sklepo-
kowane genetycznie, rozwijaj ce si w ró nym czasie. Mikromorfologi , anatomi oraz
ultrastruktur 21-dniowych zawi zków owoców odmian ‘Jonagold’ i ‘Szampion’ badano
za pomoc mikroskopii wietlnej oraz elektronowej: skaningowej i transmisyjnej. Szcze-
góln uwag zwrócono na cechy maj ce wp yw na j drno oraz trwa o owoców.
Stwierdzono, e zawi zki badanych odmian ró ni y si istotnie liczb szparek i blizn w o-
skowych obecnych na jednostce powierzchni owocu, grubo ci pok adu hipodermy oraz
jej cian komórkowych, a tak e zawarto ci depozytów zwi zków fenolowych. Na etapie
zawi zka u obydwu odmian zaobserwowano nast puj ce cechy maj ce zwi zek ze wzro-
stem lub obni eniem j drno ci i trwa o ci owoców: wosk krystaliczny w postaci p ytek,
mikrosp kania w kutykuli, szparki i blizny w oskowe oraz obecno zwi zków fenolo-
struktura, kutykula, wosk epikutykularny, mikrosp kania, zwi zki fenolowe
Accepted for print: 6.06.2014