Genetic parameters of fillet fatty acids

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 Aquaculture 556 (2022) 738292

Contents lists available at ScienceDirect

Aquaculture
journal homepage: www.elsevier.com/locate/aquaculture

Genetic parameters of fillet fatty acids and fat deposition in gilthead

seabream (Sparus aurata) using the novel 30 k Medfish SNP array
S.S. Horn a, *, M.L. Aslam a, G.F. Difford a, K. Tsakoniti b, S. Karapanagiotis b, B. Gulzari c,
J.W.M. Bastiaansen c, C. Peñaloza d, R. Houston d, B. Ruyter a, A.K. Sonesson a
a
Nofima, Norwegian Institute for Food, Fisheries and Aquaculture Research, NO-1433 Ås, Norway
b
Galaxidi Marine Farm S.A., Galaxidi 330 52, Greece
c
Wageningen University & Research, Animal Breeding and Genomics, PO Box 338, 6700 AH Wageningen, the Netherlands
d
The Roslin Institute and Royal (Dick) School of Veterinary Studies, University of Edinburgh, Midlothian EH25 9RG, United Kingdom

A R T I C L E I N F O A B S T R A C T

Keywords: Lipid-related traits are important candidates for a breeding goal for gilthead seabream, because they affect both
Omega-3 fish and human health, as well as production efficiency. However, to date there have been very few estimates of
Gilthead seabream genetic parameters for these traits, and the genetic relationship between fatty acids and other important traits
Lipid metabolism
have never been reported for gilthead seabream. Therefore, the aim of this study was to estimate genomic
Lipid deposition
Genetic parameters
heritability and genetic relationships of fat deposition traits and individual muscle fatty acids in a commercial
population of gilthead seabream using the novel ~30 k MedFish SNP array.
In total 967 gilthead seabream fed with a commercial feed were genotyped with the MedFish SNP chip which
included ~30 K informative markers for this species. On average, the fish weighed 372 g. The mean content of
eicosapentaenoic acid (EPA) + docosahexaenoic acid (DHA) was 822 mg per 100 g fillet. The heritability of
muscle fat, viscera weight and percentage viscera were in the range of 0.34–0.46. The genetic correlation of body
weight with muscle fat was 0.12, indicating that genetic variation in muscle fat is largely independent of the
weight of the fish. The heritability of the product of endogenous fatty acid synthesis (n = 240), palmitoleic acid
(16:1n-7), was high (0.43). The estimated heritability of EPA (%) and DHA (%) was 0.39 and 0.33, respectively.
Both EPA and DHA had low, non-significant genetic correlations with body weight, and DHA had a negative
genetic correlation with muscle fat (− 0.53).
It is possible to increase EPA and DHA content in gilthead seabream fillets by selective breeding. The high
heritability of 16:1n-7, a marker of de novo lipogenesis, suggests that there is a strong genetic component to this
metabolic pathway in gilthead seabream. Muscle fat deposition and body weight seem to be independent traits,
and selective breeding for faster growth is not likely to influence the proportional content of EPA and DHA.

1. Introduction Lipids are important in production of all farmed animals because

Gilthead seabream is one of the major aquaculture species in Europe,
ranked as number four, with a total production of almost 200,000 tons in
2019 (FEAP, 2020), but is at an early stage regarding selective breeding.
The first commercial breeding programs of seabream were initiated in
the early 2000's and have primarily focussed on improving growth rate
(Brown, 2004; Janssen et al., 2017; Thorland et al., 2007).
they are linked to production efficiency, health, and product quality.
Excess dietary lipids that are not deposited in the edible muscle or used
as energy source for growth are considered a loss and an indicator of low
efficiency. Constant energy excess may lead to excessive fat deposition
in and around internal organs and thereby increase the risk of metabolic
disorders, oxidative stress and inflammation (Fontana et al., 2007;
Todorčević et al., 2010).

* Corresponding author.
E-mail addresses: siri.storteig.horn@nofima.no (S.S. Horn), Luqman.Aslam@Nofima.no (M.L. Aslam), gareth.difford@nofima.no (G.F. Difford), k.tsakoniti@
galaxidimarine.farm (K. Tsakoniti), s.karapanagiotis@galaxidimarine.farm (S. Karapanagiotis), benan.gulzari@wur.nl (B. Gulzari), john.bastiaansen@wur.nl
(J.W.M. Bastiaansen), Carolina.Penaloza@roslin.ed.ac.uk (C. Peñaloza), ross.houston@roslin.ed.ac.uk (R. Houston), Bente.Ruyter@Nofima.no (B. Ruyter), anna.
sonesson@nofima.no (A.K. Sonesson).

https://doi.org/10.1016/j.aquaculture.2022.738292
Received 22 December 2021; Received in revised form 22 April 2022; Accepted 23 April 2022
Available online 27 April 2022
0044-8486/© 2022 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
S.S. Horn et al.
The lipid content of muscle tissue affects organoleptic and processing
qualities of fish fillets. In seabream, it has been shown that the taste,
colour, and juiciness of fillets are affected by the fat content (Grigorakis
et al., 2003; Grigorakis, 2007). Muscle fat also affects the nutritional
quality of fillets. Fish is an important source of the essential omega-3
long-chain polyunsaturated fatty acids (LC-PUFAs), eicosapentaenoic
acid (EPA, 20:5n-3) and docosahexaenoic acid (DHA, 22:6n-3) in human
diets. EPA and DHA have several beneficial health effects, such as pre­
venting and attenuating a range of inflammatory disorders, including
cardiovascular disease, immune dysfunction, and obesity (Calder, 2015;
Thota et al., 2018; Todorcevic and Hodson, 2015). Hence, improving the
omega-3 content of fish fillet is a point of focus in aquaculture. As a
significant part of the Mediterranean diet, seabream is an important
source of these essential and health-promoting fatty acids.
In general, omega-3 fatty acid content of fish muscle is largely
determined by the dietary fatty acid composition, which is well docu­
mented also for gilthead seabream (Ballester-Lozano et al., 2011; Gri­
gorakis et al., 2002; Izquierdo et al., 2003; Mnari et al., 2007). In
addition to feed, other factors including environmental factors (e.g.
water temperature) have also been shown to influence the omega-3 fatty
acid content of seabream muscle (Ibarz et al., 2005). Moreover, meta­
bolic factors are likely to play a role, as higher DHA level in muscle
compared to the feed have been observed in gilthead seabream
(Izquierdo et al., 2003). Omega-3 content in fish fillets can also be
regulated genetically: a recent study in gilthead seabream showed that
there is a heritable component to variation in muscle content of EPA and
DHA in gilthead seabream (Vallecillos et al., 2021). Dong et al. (2016)
estimated a heritability of 0.44 for the omega-3 levels of fillets of the
marine fish species large yellow croaker. Studies in Atlantic salmon have
revealed medium to high heritability for the omega-3 levels of fillets and
showed that genetic variation is possibly linked to omega-3 bioconver­
sion capacity (Horn et al., 2018; Leaver et al., 2011). Although marine
species have very limited capacities for omega-3 bioconversion (con­
version of the shorter chain alpha-linolenic acid (ALA) to the longer
chain EPA and DHA), some species like gilthead seabream and European
seabass do express several of the required genes. Fatty acid elongase
genes, as well as Delta-6 desaturase have been described, and are up-
regulated by dietary factors in the same manner as in salmonids (Sei­
liez et al., 2003; Tocher and Ghioni, 1999). Still, in gilthead seabream,
the omega-3 fatty acid bioconversion capacity is shown to be limited due
to a deficiency in Delta5 desaturase activity (Tocher and Ghioni, 1999).
There are, however, other metabolic factors that can influence the
omega-3 fatty acid composition in fish, such as fatty acid uptake,
deposition, and beta-oxidation (Henderson and Sargent, 1985; Horn
et al., 2019; Vegusdal et al., 2004). Although knowledge is limited in
gilthead seabream, studies have shown a preferential retention of
arachidonic acid (20:4 n-6) and DHA over EPA (Izquierdo, 1996; Kalo­
geropoulos et al., 1992; Koven et al., 2001).
The first publicly available combined-species SNP array for European
seabass and gilthead seabream, the new Axiom® MedFish SNP array,
was recently developed (Peñaloza et al., 2021). This array enables the
accurate high-throughput genotyping of ~ 30 K SNPs distributed
throughout the seabream genome. The availability of genotype data on
individuals allows the computation of genomic relationship matrix
(GRM) where the coefficient of relationships between relatives are
described more precisely with possibility of covering the deviations in
relatedness caused by Mendelian sampling (Houston et al., 2020; Niel­
sen et al., 2009). Although heritability can be underestimated or over­
estimated in influential regions with high or low linkage disequilibrium
(LD), GRM reflects more precise relatedness among individuals than the
pedigree-based relationships (Da et al., 2014). The genetic variance
components estimated using GRM are therefore possibly more precise
than pedigree-based estimates, especially when the available pedigree is
not very deep.
A key factor in the rate of genetic improvement possible for any given
trait in a breeding programme is its heritability, and its genetic
2
Aquaculture 556 (2022) 738292
correlations with other key traits in the breeding goal. Only a few studies
have reported genetic parameters of fat deposition traits in gilthead
seabream (Elalfy et al., 2021; García-Celdrán et al., 2015b; Navarro
et al., 2009b), but never using high-throughput genotyping. One study
has reported heritability of muscle fatty acids (Vallecillos et al., 2021),
but the genetic correlations between fatty acids and other important
traits have never been reported for gilthead seabream.
The aim of the current study was to estimate genomic heritability
and genetic relationships of fat deposition traits and individual muscle
fatty acids in gilthead seabream using the novel ~30 k MedFish SNP
array.
2. Material and methods
Gilthead seabream originating from the selective breeding program
of Galaxidi Marine Farm SA (Galaxidi, Greece) were used in this
experiment. All fish used in this study were the offspring of 33 males and
20 females mass spawned in a single broodstock tank over one day. The
fish were stocked in a sea cage at an average weight of 2.6 g. During the
sea phase, the fish were fed commercial feed and kept in commercial sea
cages in Galaxidi, Greece. They were slaughtered in September 2019,
after 15 months at sea and fasted prior to slaughter. From the average
body weight of ≈165 g to 270 g, the fish were fed a diet with a protein/
lipid content of 45/20, that only contained fish oil. From six to eight
months, the fish were fed a second experimental diet of a 45/16 protein/
lipid content, that included some sunflower oil. During the last two-
week phase of the trial before slaughter, the fish returned to the first
diet. Due to the logistics of sampling, slaughtering took place over
several days, therefore the number of days fasted prior to slaughter
varied from 4 to 12, which is considered in the statistical analysis of the
data.
At the time of slaughter, body weight, viscera weight, and muscle fat
measured by using Distell meter were recorded on 959 fish. Total viscera
weight was measured directly after slaughter by removing and weighing
the entire viscera, including liver and heart. The liver and heart were
also weighed separately, so for the analyses in the current study, the
viscera weight was calculated by subtracting heart weight and liver
weight from total viscera weight. Visceral fat was not measured directly,
but viscera percentage of body weight (viscera weight/body
weight*100) was used as an indicator of amount of visceral fat in the
current study (Viscera %).
2.1. Lipid measurements
Two methods of measuring muscle fat percentage were applied on
the fish studied, the non-invasive Distell fat meter (Distell (Model-FM
692, www.distell.com), and the gold standard laboratory Folch method
(MFATF). Directly after slaughter, a non-invasive fat measurement was
made on each of the 959 fish using the Distell fat meter, a handheld
microwave dielectric spectrometer. Two measurements were recorded
above the lateral line and two below, repeated on both sides of the fish.
The average of these eight measurements was used as a measure of the
total muscle fat percentage (MFATD). A calibration equation for muscle
fat in sea bream using the internal “custom calibration” setting was
generated as per the manufacturer's specifications (Distell, 2011). The
resulting calibration equation on 20 fish had a coefficient of determi­
nation of R2 = 0.79. The average of the eight measurements per fish after
use of the custom calibration was used as measure of the total muscle fat
percentage (MFATD).
The whole skinless fillet of 240 of the 959 fish were individually
homogenized, and total lipids were extracted using the Folch extraction
method (Folch et al., 1957) to record total muscle fat percentage
(MFATF). The fatty acid composition of the total lipids was determined
using the Mason & Waller method by means of methyl ester gas chro­
matography separation and flame ionization detection (Mason and
Waller, 1964). This gave fatty acid composition estimates for both
S.S. Horn et al.
proportional (% of total fatty acids) and quantitative (mg per g fillet,
based on internal standard) content of each fatty acid. For the data
analysis, outliers and error datapoints were removed through the
Interquartile Range (IQR) method. The results of the current study will
focus on the five most prevalent fatty acids of the muscle (18:1n-9, 16:0,
18:2n-6, 22:6n-3 (DHA) and 16:1n-7), as well as the omega-3 fatty acids
20:5n-3 (EPA) and 18:3n-3 (ALA).
2.2. DNA extraction and genotyping
DNA was extracted from fin clips by IdentiGEN (Dublin, Ireland),
using a crude DNA isolation method consisting of PK/Chelex extraction
followed by a dilution step. The fish were genotyped using the MedFish
array, which contains approximately 29,800 validated SNPs for gilthead
seabream (Peñaloza et al., 2021).
Genotype data were filtered using Plink software (Purcell et al.,
2007), excluding SNPs with minor allele frequency (MAF) lower than
2%, missing call rates exceeding 10%, and variants which had Hardy-
Weinberg equilibrium exact test p-value below 1e-15. In total 25,919
SNPs passed filters and quality control.
2.3. Statistical analysis
The genomic relationship matrix was generated with the GCTA
software, following the method by Yang et al. (2011), using the
following equation to estimate the genetic relationship between in­
dividuals j and k:
( )
1 ∑N xij − 2pi (xik − 2pi )
Gjk = ,
N i=1 2pi (1 − pi )
where xij is the number of copies of the reference allele for the ith SNP of
the jth individual, xik is the number of copies of the reference allele for
the ith SNP of the kth individual, and pi is the frequency of the reference
allele, estimated from the observed allele frequencies.
Variance and covariance components were estimated by residual
maximum likelihood procedures using the CGTA program. Bivariate
analyses were performed to estimate genetic correlations between traits,
using the following bivariate animal model (Henderson, 1984):
Y = XB + U + E
where Y is a matrix of phenotypic records for individuals i = 1, 2, …, n
and traits j = 1, 2, X is a matrix of the fixed effects of animal i on trait j, B
is a matrix of fxed efect solutions. U is a matrix containing the random
effects of animal i on trait j, with variance with variance G0⊗ G, where G
is the genomic relationship matrix between individuals and G0 is a ge­
netic variance–covariance matrix among traits. E is a matrix of residual
effects, that is assumed to have a variance of
[ ]
σ2e1 σ e1,e2
R = , where 1 and 2 indicate traits.
σ e1,e2 σ 2e2
Univariate analyses were performed to estimate heritability for all
traits. For the univariate analyses, matrices Y, B, U and E were reduced
to vectors and matrices G and R were reduced to scalars. Heritability
(narrow sense) was estimated as the ratio of additive genetic variance to
total phenotypic variance. Slaughter day (representing number of days
fasted) was included as a fixed effect for all traits.
Genetic coefficient of variation was determined based on the method
suggested by Burton and Devane (1953) as follows:
√̅̅̅̅̅
σ2 g
GCV = *100,
μ
Where μ is the mean of the trait, and σ 2g is the genetic variance.
As parent fish were not genotyped, it was not possible to perform
parentage assignment and develop pedigree, therefore, principal
component analysis (PCA) was performed to evaluate population
3
Aquaculture 556 (2022) 738292
structure and get an idea on number of clusters possibly representing full
and/or half-sib families. The PCA based cluster analysis showed that the
population is quite homogenous (little to no cluster differentiation),
with relatively low variation explained by first two PCAs, 4.9% and
3.8% variation explained by PCA1 and PCA2, respectively (Fig. 1). The
cluster analysis showed ~42 clusters with number of individuals
ranging from 2 to 53 individuals per cluster with a mean cluster size of
23 individuals.
3. Results and discussion
The body size and muscle fat level were within the normal range for
commercially produced harvest sized gilthead seabream, with a mean
body weight of 372 g, ranging from 124 to 546 g (Table 1). The coef­
ficient of variation for all production traits surpassed 0.15, showing that
there was substantial individual variation in these traits.
3.1. Genetic parameters of lipid related production traits
The heritability of muscle fat, viscera weight and viscera % were all
above 0.3, showing that there is a substantial genetic component to
these traits (see Table 2). Viscera % had an especially high heritability
estimate of 0.46, which was almost equal to the estimated heritability of
viscera weight with body weight included as covariate in the statistical
model (0.45 ± 0.06, result not shown). This result is in line with the
previously reported heritability of 0.5 for visceral fat in gilthead seab­
ream by Navarro et al. (2009b), although García-Celdrán et al. (2015a)
reported a heritability of 0.2 for visceral fat with body weight as a co­
variate in the model. In the current study, the observed variation in
viscera weight was assumed to mainly be due to the size of the fish and
the amount of lipids stored here; hence the viscera weight expressed as
% of body weight was used as an indicator of amount of visceral fat. The
genetic correlation between body weight and viscera weight was 0.72,
supporting that there is some genetic variation in viscera weight that is
independent of the weight of the fish, which could be variation in dis­
tribution of lipid deposition.
Previously reported heritability of muscle fat ranges from 0.05 to
0.31 in gilthead seabream (Elalfy et al., 2021; García-Celdrán et al.,
2015a; Navarro et al., 2009b; Vallecillos et al., 2021). In the current
study, both methods of recording muscle fat showed heritability slightly
higher than previous estimates: MFATD had a heritability of 0.46, and
MFATF of 0.34. The considerably smaller dataset is reflected in the
larger standard error for MFATF (Table 2). The differences in heritability
estimates between studies could be partly related to the accuracy of the
recording technique. Moreover, there are differences in the statistical
models used. García-Celdrán et al. (2015a) and Vallecillos et al. (2021)
included body weight as a covariate. In the current study, body weight
was not included as a covariate in the statistical model, but when tested,
body weight as covariate did not have a significant effect on heritability
estimates of the muscle fat traits (results not shown). This is due to that
the genetic correlation between body weight and muscle fat was very
low (rg = 0.12 for MFATD and rg = 0.13 for MFATF), which was in
accordance with the low phenotypic correlations observed (Fig. 2 &
Table 3). A low genetic correlation between body weight and muscle fat
is in agreement with the estimate reported by Navarro et al. (2009a) (rg
= 0.12). However, estimates reported by García-Celdrán et al. (2015b)
and Vallecillos et al. (2021) are higher (0.29 and 0.59, respectively).
This could explain the differences in statistical models and heritability
estimates. Furthermore, these differences in estimates of heritability and
genetic correlations may be due to genetic differences among the studied
populations, as well as genotype-by-environment interactions (Gulzari
et al., 2022; Kause et al., 2002).
The low genetic correlation between body weight and muscle fat
observed in the current study of gilthead seabream is contrary to find­
ings of salmonid species such as Atlantic salmon where muscle fat per­
centage is highly genetically correlated with body weight, i.e. rg =
S.S. Horn et al. Aquaculture 556 (2022) 738292

Fig. 1. PCA plot of the population structure.

Table 1
Descriptive statistics for production traits.
N Mean (SD) Min Max CV (%)

Body weight (g) 967 372 (64) 124 546 17

Viscera weight (g) 911 22.9 (6.0) 5.4 39.6 26
Viscera % 911 6.1 (1.1) 3.3 9.8 18
MFATD (%) 960 7.6 (1.54) 2.8 11.9 18
MFATF (%)* 238 9.4 (0.1) 3.9 14.8 20

SD = standard deviation. CV = coefficient of variation. Viscera % = (viscera

weight/body weight), MFATD = muscle fat % by Distell meter, MFATF = muscle
fat % by Folch chemical analysis.

Table 2
Heritability (h2) of production traits.
Trait N Va Ve h2 GCV %

Body weight 967 1435 (271) 2842 (187) 0.34 (0.05) 10

Viscera weight 911 14.87 (2.73) 23.31 (1.7) 0.39 (0.06) 17
Viscera % 911 0.58 (0.09) 0.67 (0.05) 0.46 (0.05) 12
MFATD 960 1.03 (0.16) 1.21 (0.09) 0.46 (0.05) 13
MFATF 238 1.13 (0.46) 2.2 (0.38) 0.34 (0.12) 11

Standard errors in brackets. Va: Additive genetic variance. Ve: Residual vari­
ance. GCV = Genetic coefficient of variation. Viscera % = (viscera weight/body
weight), MFATD = muscle fat % by Distell meter, MFATF = muscle fat % by Folch
chemical analysis. Fig. 2. Scatter plot of body weight against muscle fat, including the regression
line and correlation coefficient, n = 238.
0.45–0.84 (Powell et al., 2008; Tsai et al., 2015; Vieira et al., 2007).
There are similar findings in the salmonids Rainbow trout (rg =
0.30–0.52) (Blay et al., 2021) and European white fish (rg = 0.62–0.64) Table 3
Genetic (upper triangle) and phenotypic (bottom triangle) correlations.
(Janhunen et al., 2017). This difference is likely explained by the
inherent differences in the mechanisms of lipid deposition between the Body Viscera Viscera % MFATD MFATF
fish species. For example, in Atlantic salmon, the muscle is a main lipid weight weight

storage site, while in gilthead seabream very little fat is stored in muscle, Body 0.72 (0.06) 0.25 0.12 0.13
and the viscera is the main lipid storage site (McClelland et al., 1995). weight (0.12) (0.12) (0.20)*
Viscera 0.74 0.86 0.20 0.19
These differences in lipid storage are likely due to differences in
weight (0.03) (0.12) (0.20)*
behavioural adaptation to their natural habitats; seabream is a bottom- Viscera % 0.22 0.81 0.23 0.11
dwelling sedentary fish species, while Atlantic salmon is an active sur­ (0.11) (0.20)*
face feeder species, and needs energy available in their locomotory MFATD 0.28 0.24 0.18 0.82
muscles (McClelland et al., 1995). (0.09)*
MFATF 0.15** 0.21 0.18 0.60
There was a low genetic correlation between the two fat deposition
traits MFATD and Viscera % (rg = 0.23). This agrees with Navarro et al. Standard errors in brackets.
*
(2009a), who found an even lower correlation of 0.05, and used a direct Only converged with no covariates in the statistical model.
**
measure of visceral fat - manual removing and weighing of visceral fat p > 0.01.

4
S.S. Horn et al. Aquaculture 556 (2022) 738292

deposits. This shows that deposition of fat in these two lipid deposits with body weight (Table 6). However, DHA (but not EPA) was signifi­
(muscle and viscera) are genetically different traits, similar to what has cantly correlated with muscle fat, which is in agreement with Vallecillos
been described in Rainbow trout (Kause et al., 2007). et al. (2021). Thus, including muscle fat as a covariate in the model
The genetic correlation between the two methods of measuring would likely reduce the heritability estimate for DHA in this data
muscle fat was high; 0.82 (Table 3), but lower than reported by a recent material.
study on European seabass, where the genetic correlation between the One limitation of this study is the lack of pedigree, which may have
same two methods on the same trait was close to unity at 0.96 ± 0.03 biased the estimation of genetic parameters, as population stratification
(Difford et al., 2021). Pearson's correlation coefficient for the two can inflate estimates of genetic relatedness (Dandine-Roulland et al.,
methods was 0.6, showing that the Distell meter was not highly accurate 2016). However, the evaluation of population structure by principal
for phenotypic recording of muscle fat percentage in this dataset. component analysis showed that the population is quite homogenous
with little to no cluster differentiation (Fig. 1). Also, previous studies
3.2. Muscle fatty acid composition have shown that genomic information can more accurately reflect the
relationships between individuals than pedigree information (Da et al.,
The fatty acid composition of muscle was recorded as both propor­ 2014; Wang and Da, 2014). The increased accuracy of genomic pre­
tional content (% of total fatty acids) and quantitative content (mg/g diction compared with pedigree prediction is evident in a range of
tissue) on 240 fish (Table 4). The phenotypic variation in proportional aquaculture species (reviewed in Houston et al. (2020)).
content was low, while the variation in quantitative content was higher, The high heritability for both EPA and DHA estimated in the current
reflecting the variation in total muscle fat content. The major fatty acids study demonstrate that selective breeding is a promising tool for
in the muscle, constituting more than 60% of muscle fatty acids, were increasing muscle content of these essential nutrients in the fillet of this
oleic acid (18:1n-9), palmitic acid (16:0), and linoleic acid (18:2n-6). species. This agrees with studies in Atlantic salmon that have shown
Oleic and linoleic acid are the major fatty acids from plant oils in fish significant heritability of EPA and DHA content of muscle (Leaver et al.,
feed, while palmitic acid is the most common saturated fatty acid found 2011; Horn et al., 2018). Wang et al. (2019) also showed that there is a
in all animals. The mean proportional content of EPA and DHA in the genetic component to omega-3 content of Asian seabass fillets. Herita­
gilthead seabream muscle was 2.43% and 6.95% of total lipids, bility estimates of DHA in tilapia and common carp have been close to
respectively. The mean quantitative content of EPA and DHA was 2.14 zero, which could be due to the very low quantities of DHA detected
and 6.08 mg/g, respectively, i.e. 822 mg per 100 g fillet in total. (Nguyen et al., 2010; Prchal et al., 2018). The estimates for both fish
Considering the recommendations on daily intake of omega-3 FAs for species indicated that EPA had a considerably higher heritability than
the general human public of 250–500 mg EPA + DHA (Cunnane, 2004; DHA, while in Atlantic salmon, the heritability of EPA was lower than
GOED, 2014), farmed seabream is a good source of the essential nutri­ DHA (Horn et al., 2018).
ents EPA and DHA. Alpha-linolenic acid (18:3n-3; ALA) had the lowest heritability of all
fatty acids (0.03). The ratio of DHA/ALA was included in the analysis as
an indicator-trait for the conversion of ALA to DHA (omega-3 biocon­
3.3. Heritability and genetic correlations of fatty acids in gilthead
version), but is in this case likely reflecting variation in DHA due to the
seabream fillets
very low variation in ALA. This low variation, together with the low
general content of ALA in the muscle of gilthead seabream, indicates
The estimated heritability of the proportional content of both EPA
that omega-3 bioconversion of ALA to DHA is not substantial, and not
and DHA was relatively high, at 0.39 and 0.33, respectively (Table 5).
very important for the overall DHA content in the muscle of this species.
These estimates are substantially higher than those recently reported by
This implies that the phenotypic and genetic variation observed in EPA
Vallecillos et al. (2021), at 0.05 for EPA and 0.11 for DHA. This differ­
and DHA levels are linked to other metabolic factors, such as deposition
ence is likely due to the different statistical models used, as Vallecillos
or beta-oxidation, rather than omega-3 bioconversion. This is in accor­
et al. (2021) used a Bayesian model, which is quite conservative, and
dance with a study reporting that the omega-3 bioconversion capacity is
included body weight as a covariate in the model, which could remove
limited in gilthead seabream due to a deficiency in Delta5 desaturase
some of the genetic variation in the fatty acid traits. In the current study,
activity, catalyzing conversion of 20:4n-3 to EPA (Tocher and Ghioni,
we chose not to include body weight as a covariate for fatty acids, as
1999).
both % EPA and % DHA had low and non-significant genetic correlations
The estimates of genetic correlations showed that the proportional
content of each fatty acid was differently correlated with muscle fat
Table 4 (Table 6). The standard errors on the estimates were large due to the
Descriptive statistics of proportional (% of total fatty acids) and quantitative
small dataset on fatty acids, but the genetic correlations agreed with the
(mg/g tissue) content of the top five most prevalent fatty acids in muscle, and the
phenotypic correlations. The fatty acids with the strongest positive
omega-3 fatty acids EPA (20:5n-3) and ALA (18:3n-3).
correlations with muscle fat were 18:1n-9 and 16:1n-7 (Table 6), both of
Fatty Proportional content (%) Quantitative content (mg/g) which are products of de novo lipogenesis, synthesis of fatty acids,
acid
Mean Min Max CV Mean Min Max CV although 18:1n-9 is also found in the feed. This indicates that a genetic
(SD) (%) (SD) (%) predisposition for higher muscle fat is due to a higher inherent de novo
18:1 n-9 29.0 27.4 30.6 2 25.6 12.5 39.3 20 lipogenesis activity. The high heritability of % 16:1n-7, which is almost
(0.05) (0.33) exclusively a product of de novo lipogenesis, also indicates that the rate
16:0 15.7 14.4 16.9 3 13.8 7.1 21.2 19 of de novo lipogenesis in gilthead seabream muscle is a heritable trait
(0.03) (0.17)
18:2 n-6 15.6 13.9 17.3 4 13.8 7.1 21 19
(Table 5).
(0.04) (0.17) The genetic correlations indicated that the observed genetic varia­
22:6 n-3 7.0 6.1 7.9 6 6.1 3.5 8.5 16 tion in % EPA and % DHA is independent of body weight, as both % EPA
(DHA) (0.03) (0.06) and % DHA had low and non-significant genetic correlations with body
16:1 n-7 5.1 4.4 5.9 6 4.5 2.1 7 22
weight (Table 6). The proportional content of EPA had a weak positive
(0.02) (0.06)
20:5 n-3 2.4 2 2.9 7 2.1 1.1 3.2 20 (non-significant) correlation with muscle fat, while DHA had a negative
(EPA) (0.01) (0.03) genetic correlation. This difference between EPA and DHA likely reflects
18:3 n-3 2.4 2.2 2.6 3 2.1 1.1 3.2 20 the metabolic differences between the two fatty acids, and their roles in
(ALA) (0.01) (0.03) muscle tissue (Calder, 2006; Stillwell and Wassall, 2003). The observed
N = 240. CV = coefficient of variation. SD = standard deviation. relationship between DHA and muscle fat is seen in several species and

5
S.S. Horn et al. Aquaculture 556 (2022) 738292

Table 5
Heritability (h2) of fatty acid traits.
Trait % mg/g
2
Va Ve h GCV (%) Va Ve h2 GCV (%)

16:0 0.06 (0.03) 0.16 (0.02) 0.28 (0.11) 1.57 2.17 (0.84) 4.21 (0.70) 0.33 (0.11) 10.67
16:1n-7 0.04 (0.01) 0.05 (0.01) 0.43 (0.12) 3.75 0.34 (0.12) 0.56 (0.10) 0.38 (0.12) 12.88
18:1n-9 0.16 (0.07) 0.32 (0.06) 0.34 (0.12) 1.40 9.25 (3.33) 15.65 (2.68) 0.36 (0.11) 11.88
18:2n-6 0.13 (0.06) 0.32 (0.05) 0.28 (0.12) 2.27 2.07 (0.78) 4.27 (0.66) 0.33 (0.11) 10.44
ALA 0.00 (0.00) 0.01 (0.00) 0.03 (0.08) 0. 59 0.06 (0.02) 0.10 (0.02) 0.35 (0.11) 11.40
EPA 0.01 (0.00) 0.02 (0.00) 0.39 (0.11) 4.28 0.07 (0.02) 0.12 (0.02) 0.37 (0.12) 12.56
DHA 0.05 (0.02) 0.10 (0.02) 0.33 (0.12) 3.14 0.32 (0.13) 0.61 (0.10) 0.35 (0.12) 9.21
DHA/ALA 0.01 (0.00) 0.03 (0.00) 0.25 (0.11) 3.40 – – – –

% = proportional content of fatty acids, mg/g = quantitative content of fatty acids, Va: Additive genetic variance. Ve: Residual variance. Standard errors in brackets.
GCV = genetic coefficient of variation. N = 240.

correlations indicated that increasing muscle fat would actually reduce

Table 6
the proportional content of DHA in the muscle, leading to a less desir­
Phenotypic (rP) and genetic (rG) correlations between production traits and fatty
able fatty acid profile of the seabream fillet (Table 6). On the other hand,
acids.
the genetic correlations between proportional contents of muscle fatty
Trait rP rG (SE)
acids showed a potential to select for an overall healthier fatty acid
BW * % EPA 0.24 0.17 (0.18) profile in muscle (Table 6). DHA was positively correlated with EPA (rg
BW * % DHA 0.08** − 0.03 (0.20) = 0.55), and negatively correlated with the main saturated fatty acid
MFATF * % EPA − 0.05** 0.11 (0.25)
(16:0) and the main omega-6 fatty acid (18:2n-6) in muscle (rg = − 0.48
MFATF * % DHA − 0.56 − 0.53 (0.20)
MFATF * % 18:1n-9 0.5 0.81 (0.17) and − 0.37, respectively). Thus, according to these results, selection for
MFATF * % 16:1n-7 0.4 0.51 (0.20) increased proportional content of DHA will result in a fish fillet with a
MFATF * 18:2n-6 − 0.07** − 0.20 (0.29) higher proportion of marine omega-3 fatty acids, and lower proportion
MFATF * 16:0 0** − 0.15 (0.30)
of saturated fatty acids and pro-inflammatory omega-6 fatty acids.
DHA * EPA 0.42 0.55 (0.19)
DHA * 18:2n-6 − 0.22 − 0.37 (0.29)
However, a lower selection response may be expected using this strategy
DHA * 16:0 − 0.23 − 0.48 (0.25) as the genetic coefficients of variation was lower for proportional
DHA * 16:1n-7 − 0.04** 0.05 (0.26) compared to quantitative content of EPA and DHA (Table 5).
DHA * 18:3n-3 − 0.22 nc Even though both quantitative and proportional content of EPA and
DHA * 18:1n-9 − 0.63 nc
DHA are highly heritable, getting genetic response in fatty acid traits is
MFATF = Muscle fat % by Folch chemical analysis, BW = body weight. SE = not straight forward. With low phenotypic and genetic coefficients of
standard error. EPA = 20:5n-3. DHA = 22:6n-3. Nc = no convergence. ** = p > variation it is difficult to get a reasonable selection response, even when
0.01. heritability is high. We recommend future research into economic
weights and selection indices which are outside the scope of this paper.
tissues and can be explained by DHA being a PUFA with important roles Another challenge with implementing selection for fatty acid traits is
in cell membranes that is therefore found in phospholipids (PL) to a that phenotyping requires costly and time-consuming chemical ana­
larger degree than in triglycerides (TG). When muscle fat increases, the lyses, which make large-scale data recording challenging. New rapid
proportion of PL relative to TG decreases, and therefore the % DHA methods for predicting fatty acid composition in fish fillet are being
decreases (Tocher, 2003). developed, such as Raman and near infrared spectroscopy (e.g. EWOS
SalmoNIR technology from Cargill) (Afseth et al., 2006; Bekhit et al.,
3.4. Selection strategies for increased omega-3 fatty acid content of fillets 2014; Blay et al., 2021), which can make practical implementation of
selection for these traits more feasible.
The heritability of EPA and DHA indicated that the content of
healthy omega-3 fatty acids in seabream fillets can be increased through 4. Conclusions
selective breeding, and there are two possible strategies to achieve this:
by increasing the quantitative content (mg per g muscle), or by In the gilthead seabream, muscle fat deposition and body weight
increasing the proportional content (% of total muscle fatty acids). seem to be independent traits, and selective breeding for faster growth is
As muscle fat percentage increases, so does the quantitative content not likely to influence the proportional content of EPA and DHA. The
of all fatty acids. Therefore, the heritability of the quantitative content of fatty acid 16:1n-7, a marker of de novo lipogenesis, had a high herita­
all fatty acids mirrored the heritability of muscle fat and were all close to bility (0.43), indicating that there is a strong genetic component to this
0.34 (Table 5). This was confirmed in the estimates of genetic correla­ metabolic pathway in seabream.
tions, where the quantitative content of all fatty acids had a genetic It is possible to increase EPA and DHA content in gilthead seabream
correlation to muscle fat close to unity (results not shown). This implies fillets by selective breeding, as the estimated heritability of EPA (%) and
that an increase in the quantitative content of EPA and DHA can be DHA (%) was 0.39 and 0.33, respectively, and there was a positive ge­
achieved by selective breeding for increased muscle fat content, which netic correlation between the two fatty acids (0.55). There is a potential
could be achieved by e.g. the Distell fat meter for non-invasive pheno­ to select for a healthier overall fatty acid profile in muscle of seabream
typic recording. It should then be considered that increased muscle fat by selection for increased proportional content of DHA, as DHA was
would influence the organoleptic qualities of the fish (Grigorakis, 2007). negatively correlated with the major omega-6 fatty acid (18:2n-6) and
It should also be considered that selective breeding for increased muscle the major saturated fatty acid (16:0).
fat content would result in an increase in all fatty acids, including the
less desirable omega-6 fatty acids. This is critical from a nutritional CRediT authorship contribution statement
health perspective, as it is not only the amount of EPA and DHA that is
important, but also the ratio of omega-3 to omega-6 fatty acids (Saini S.S. Horn: Methodology, Writing – original draft, Visualization,
and Keum, 2018; Simopoulos, 2002). In addition, the genetic Writing – review & editing, Formal analysis. M.L. Aslam: Methodology,

6
S.S. Horn et al.
Writing – review & editing, Formal analysis. G.F. Difford: Methodology,
Writing – review & editing, Formal analysis. K. Tsakoniti: Resources,
Conceptualization, Data curation. S. Karapanagiotis: Resources, Data
curation, Writing – review & editing. B. Gulzari: Writing – review &
editing. J.W.M. Bastiaansen: Writing – review & editing. C. Peñaloza:
Resources, Writing – review & editing. R. Houston: Resources, Writing
– review & editing. B. Ruyter: Data curation, Conceptualization,
Writing – review & editing. A.K. Sonesson: Funding acquisition, Project
administration, Supervision, Conceptualization, Methodology, Writing –
review & editing.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
This study was made possible by the EU project MedAID (H2020
grant agreement No 727315). Authors would like to thank Galaxidi
Marine Farm S.A. for their participation in the study and its staff for
providing necessary help and facilities for raising the population and the
experimental data collection. Authors are grateful to Målfrid Tofteberg
Bjerke and Anne Marie Langseter for skilful technical assistance.
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Another random document with
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history that the warship was attended to; we may even go so far as
to assert that it was not until the time of the Middle Ages that the
merchant ship both of the Mediterranean and the North of Europe,
made any real progress. As long as civilisation was scanty and
pirates were rampant on every sea, commerce was bound to remain
at a standstill. Indeed, in the time of the early Greeks, it was thought
no act of discourtesy to ask a seafaring stranger whether he was a
pirate or merchant. So accustomed are we in these days to peace
and plenty that we have need to remind ourselves constantly that
there were no trade routes kept open, no policing of the seas, no
international treaties nor diplomatic relations to prevent a peaceful
merchant ship from disappearing altogether on the high seas, or
staggering into port with the loss of her cargo and most of her crew.
The Egyptian stern still survives in these mosaics with
modifications, but the greatest difficulty the naval architects appear
to have had was with the bow. What to do with the ram-like entrance
has obviously been a source of great worry. In the end, so that the
merchant ship might not look too war-like, a curve has been added
above the bulwarks at the bows to balance the curve at the waterline
of the ram. The rowing arrangements exhibit a square hole in the
gunwale for the oar to pass through.
Of the sailing boats and ships depicted in mosaic the corbita
shows a freer design than the others. She is more or less crescent-
shaped and not unlike the earlier caravels in hull. A ship of burthen,
she has a mast, and the steering oar is seen at the starboard.
Another illustration of this type of “corvette” is shown with a steering
oar at each side, the sail furled to the yard, a couple of braces and
the mast supported by six shrouds—three forward and three aft. The
mast has a great rake forward, and there appears to be a narrow
platform running round the hull as a side-walk, a relic, no doubt, of
the flying deck that kept the marines separate from the rowers.
Another sailing ship called the catascopiscus obviously derived
her name from the corresponding Greek word meaning to
reconnoitre or scout; for she was famous as a light, fast-sailing ship.
Her mast and sail are shown in the mosaic, as well as the halyards
and the brailing lines.
 The actuaria was a light, easily propelled ship, similar to the last.
The mosaic (reproduced in Fig. 24) shows the sail furled to the yard
and, what is significant, a rope-ladder, up which one of the sailors is
ascending. Of the other two men one is sculling with two oars, while
the captain is seen in the bows holding a mallet, which he knocks on
the boat that the sculler may keep correct time and rhythm in a
manner not very far separated from the exhortations of the “cox” of
our University eights. This was the kind of ship which Cæsar
employed during an expedition to Brittany, and will be referred to
again in the next chapter.

Fig. 24. Navis Actuaria.

From a mosaic at Althiburus, near Tunis.

Another sailing ship, called by the artist a myoparo, shows two

halyards, and the sail divided curtain-like as we saw in the
Phœnician ships. She also has the Egyptian stern and a modified
galley bow. The myoparo was a light, swift vessel, chiefly used by
pirates. The stem of the English word “peir” (meaning to attempt to
rob) is thus found in the name of the ship. Plutarch makes use of the
name of this species of ship. The prosumia contains just such a sail
as we saw in Fig. 21, the brails being very clearly shown. A sailing
ship called a ponto has a small artemon foremast and main. The
former has shrouds to support it, but the yard and sail are not shown.
They would be kept in the hold somewhere, and only fitted when
specially needed. This ship is of Gallic origin, and is mentioned by
Cæsar, who refers to the “pontones quod est genus navium
Gallicarum.”[22] Finally, in these mosaics, we have the cladivata, a
ship that resembles the vessel referred to by Mr. Torr in his “Ancient
Ships” as having been found at Utica, and belonging to about the
year 200 a.d. This cladivata has also two masts and sails of similar
size, with the brailing arrangement of this period as already shown.
There is some uncertainty concerning the derivation of the word, but
it possibly owes its origin to being named after Claudius.
Such, then, was the development of the sailing ship in the waters
of Southern Europe. We shall now, leaving behind the first ships that
sailed the Mediterranean, proceed in our enquiry to the shores of
Northern Europe, and consider what was the nature of their ships
which had to voyage under conditions far less encouraging than
those of the warm southern seas.
 C H A P T E R I V.
T H E E A R LY S H I P S O F N O R T H E R N E U R O P E .

he evidence that we possess, in order accurately

to judge, of the early ships that sailed the seas of
the Baltic, German Ocean, Bay of Biscay, and
English Channel, is both conclusive and diverse.
We have in the writings of Cæsar and Tacitus,
many details of ships that are of considerable
interest. This literature is supplemented by the old
Sagas[23] of Scandinavia, which, though highly
informative, err on the side of exaggeration. Rock sculptures existing
in the land of the Vikings, though somewhat the subject of
controversy, are, in the writer’s opinion, of real, valuable help in the
study of sailing ships. There is also some evidence of later ships in
the old coins of Northern Europe. But it is when we come to the
important excavations that have revealed—nearly always
accidentally—the ships of a bygone age, many hundreds of years
old, that we are confronted with the most undeniable and complete
source of information that one could desire.
These excavations have revealed discoveries of two kinds, which
we shall deal with as we proceed. In Great Britain, and in Germany,
various examples of the prehistoric “dug-out” have been unearthed.
The Museums of Edinburgh, York, Bremen, and Kiel, happily contain
these interesting craft, preserved for the wonder of future
generations. The second class is more valuable still, and far more
picturesque, for thanks to the burial customs of the old sea-chiefs,
there have been excavated from certain mounds in Norway,
wonderful old Viking ships in a state of preservation that is
remarkable when we consider how many centuries they have lain
under the earth. Therefore, fortunate as we deemed ourselves in
being able to have such sources of information as models and
reproductions of the ships of Southern Europe, we are far more
happy in our present section for we can go to the fountain head
direct—the ships themselves.
To us members of the Anglo-Saxon race, the importance of the
forces at work during the period we are about to consider cannot be
lightly estimated. The influence of the Viking, or double-ended type
of ship, dominated the whole of the coast-line from Norway to the
land as far south as the northern shores of Spain, right from the
period that followed the construction of mere dug-outs, until almost
the close of the fifteenth century of our era. That is to say, as soon as
ever the North European became sufficiently civilised to build rather
than to hew his craft: as soon as he undertook the making of ships
rather than of boats—he came under the power of that naval
architecture, which we see illustrated in the ships of the Veneti and
Scandinavians; and, irrespective of geographical position, of
language, of tribe or of nation, the civilised inhabitants living on that
vast stretch of littoral, from the North Cape to the southern boundary
of the Bay of Biscay, continued in the same conventions of design
and build for many hundreds of years. It is a striking proof of the
accurate knowledge in shipbuilding and ship-designing possessed
by these early Northerners when we remember that, even to this
day, that influence, far from disappearing, shows a strong tendency
to increase, at any rate, in the architecture of yachts and fishing
boats. Thus, the Egyptians, who moulded for ever afterwards the
lines of the ships of the Mediterranean, have in Northern Europe,
their counterpart in the Norsemen. What the galley was to the south,
the Viking ship has been to us living in colder climes.
The obvious question occurs at this point as to what, if any, is the
connection between the Mediterranean galley and the ship of the
North Sea. That there is some similarity will be realised when we
collect the following characteristics. And first, the very shape of both
kinds of vessels—long, narrow, flat-bottomed; then the arrangement
of the large squaresail with its braces and rigging; the mode of
steering at the side; the pavisado that ran round the ship to protect
the men from the enemy; the spur with which they rammed the
enemy’s ships; the girdle that went round the ship to prevent
damage caused by ramming; the ornamentation of the head of the
vessel; their very methods of naval warfare, and finally, their
adoption later of fore-castles and stern-castles—what else do these
similarities show but that there existed a common influence? With
such evidence before us, it becomes somewhat difficult to find
agreement with those who contend that between the two classes of
ships there is no connection whatever, except such as chance might
have brought about. I am not denying that there are important
differences between the ships of the two seas, but I contend that
such important resemblances to each other need an explanation
more scientific than can be ascribed to chance.
But assuming that we are right in our surmise, by what means
were these early Norwegians affected by the southern design? Were
they influenced by Roman civilisation? That they certainly were not.
Then the southerners came to them?[24] Here is our contention.
Though we have no actual proof, it seems justifiable to suppose that
those great travellers and sea-folk, the Phœnicians, who, we have
seen, were unsurpassed in their time for seamanship and
shipbuilding; who have been said to have voyaged to the setting sun
as far as America, and to have crossed the Bay of Biscay to Ireland
and Cornwall, might have taken advantage of the prevailing westerly
wind which blows across our land and have held on until they had
touched the shores of Denmark or Norway. But why should they, do
you ask? We have seen that the Phœnicians were not merely great
sailormen, not merely adventurous, not merely eager explorers, but
practical business-men, merchants, traders. If they had found ore in
Cornwall, would they not have been inclined to seek other lands for
what they could barter or wrest ere returning to their own homes?
Even supposing the Phœnicians never crossed to South America,
we know that they circumnavigated Africa. A land that bred seamen
of that daring and ability would not be lacking in the kind of men to
discover Norway.
And there is still another reason, it seems to me, why the
Phœnicians might have felt tempted to go eastward after Cornwall.
Ignorant as they were of the world’s geography, might they not have
thought that, just as by sailing round always to the starboard they
had encircled Africa, so having performed roughly a semi-circle from
the Mediterranean to the English Channel, if they kept their course
over the wilderness of sea in front of them, they would ultimately find
that Europe, like Africa, was an island, too, and that the nearer they
approached the rising of the sun the sooner would they see their
homes again?
And if we are told to explain the differences between, on the one
hand, the ships of the Phœnicians or their later descendants the
Greeks and Romans, and on the other, those of the Vikings, it is but
natural that, given a general design which has originated in the
smoother waters of the Mediterranean, it must necessarily be
somewhat modified for the nasty seas of the Baltic and German
Ocean, where sudden changes of wind are but the harbingers of the
rapid approach of bad weather. Cæsar, when he came north into
Brittany, was struck, in comparing the ships of the Veneti with his
own, by the superior seaworthiness of the former, and adds
significantly that “considering the nature of the place and the
violence of the storms, they were more suitable and better
adapted.”[25] There is to-day a far greater difference in England
between the sailing ships of one port and another than there was
between the old Viking vessels and those of the Phœnicians. If you
cruise round from one coast of Great Britain to another, you will find
in the Scotch fishing craft, the Yorkshire cobble, the Yarmouth fishing
smack, the Lowestoft “drifter,” the Thames “bawley,” the Deal galley,
the Itchen Ferry transom-sterned cutters, the Brixham “Mumble
Bees,” the Falmouth Quay-punt, the Bristol Channel pilot, and the
Manx lugger, a wonderful complexity of designs and rigs, but the
reason is always that that particular design and rig have been found
to be the most suitable adaptation for each particular coast.
So it was with the Vikings. They modified the Phœnician design to
their local requirements, without, nevertheless, neglecting those
features essential to a good ship. After they had been shipbuilders
for some time they would rapidly learn for themselves the values of
length and beam, of draught and sweet lines, of straight keel, with
high stem to breast a wave, and high stern to repel a following sea.
Double-ended as they were, there was a reason for this essential
difference from the Phœnicians. Such seas as they had in the North
would not suffer their ships to be beached always in fine weather. So
in order that they could be brought to land with either end on, and in
order, too, that in sea-fights they might easily manœuvre astern or
ahead, the Viking ships were built with a bow both forward and aft.
But long before ever the Phœnician ships came to the shores of
Northern Europe there were boats and sailing ships. No doubt the
prehistoric man in the north was driven to finding some means of
transportation across the fjord by the same stern mother Necessity
that first induced that primitive whom we saw learning his elementary
seamanship on the Tigris or Euphrates. That ancient Northerner of
the Stone Age made a wonderfully historic discovery when he found
that he could make an edge to his stone, and that thereby he was
able to cut both flesh and wood. “For,” says Mr. Eiríkr Magnússon in
his interesting essay,[26] “on the edge, ever since its discovery, has
depended and probably will depend to the end of time the whole
artistic and artificial environment of human existence, in all its
infinitely varied complexity.... By this discovery was broken down a
wall that for untold ages had dammed up a stagnant, unprogressive
past, and through the breach were let loose all the potentialities of
the future civilisation of mankind. It was entirely due to the discovery
of the edge that man was enabled, in the course of time, to invent
the art of shipbuilding.”
The monoxylon—the boat made from one piece of timber—as
fashioned by the early sailorman of the Stone Age, is even still used
in parts of Sweden and Norway. Indeed it still bears the name which
is the equivalent of “oakie,” showing that it was originally made out of
the oak-trunk, which is the thickest and therefore the most suitable
trunk to be found in the forests of the North Sea coast, a region, that
in the time of the Stone Age was densely wooded with oak trees.
Afterwards, this monoxylon or dug-out, in order that she may be
made so strong as to carry as many as forty men, is strengthened
with ribs, and the flat bottom has the modification of a keel added.
The vessel that was found at Brigg in Lincolnshire in May 1886 (see
Fig. 25) is of this kind. A similar kind of boat was found in the
Valdermoor marsh in Schleswig-Holstein during the year 1878, and
is now in the Kiel Museum. As there are other similar boats in
existence, perhaps it may interest the reader if we deal with these
discoveries a little more fully.
The Valdermoor boat has the following dimensions: length 41 feet,
greatest width 4.33 feet, depth inside 19 inches, depth outside 20½
inches. The thickness of the wood is 1½ inches at the bottom and
1¼ at the top. The boat had eleven ribs, of which nine now exist. On
the gunwale between the ribs, eleven holes were made for inserting
oars. Both the stem and stern are sharp. The keel, 6½ feet in length,
is worked out of the wood at both ends of the boat, leaving the
middle flat. I am sorry not to be able to present an illustration of this
before the reader, but the director of the Kiel Museum informs me
that the boat is in such a position as to prevent it being
photographed.
However, the Brigg boat is very similar to the Valdermoor and may
serve the purposes of illustration equally well. This craft was found
by workmen excavating for a new gasometer upon the banks of the
river Ancholme, in North Lincolnshire. It had been resting apparently
on the clay bottom of the sloping beach of an old lagoon. It was
obviously made out of the trunk of a tree, and perfectly straight, its
dimensions being: 48 feet 6 inches long, about 6 feet wide, 2 feet 9
inches deep. The stern represents the butt end of a tree with
diameter of 5 feet 3 inches. The cubic contents of the boat would be
about 700 cubic feet. The prow is rounded off as if intended for a
ram, and a cavity in the head of the prow appears to have been
intended for a bowsprit, whereby the forestay could be made fast. In
fact, a piece of crooked oak suitable for this purpose was found
adjacent to the prow. Whilst the bottom of this dug-out is flat, the
sides are perpendicular and there is a kind of overhanging counter at
the stern.
The boat was formerly in the possession of Mr. V. Cary-Elwes,
F.S.A.,[27] to whom I have to express my thanks for his courtesy in
supplying me with some information regarding the boat here
reproduced. The ship was offered by this gentleman to the British
Museum, but was declined as being too big. It therefore remains in a
small provincial town difficult of access and for the most part
unknown. It would be impossible to remove the craft now, without
risk of total destruction, but is it not a little humiliating that continental
and provincial museums should see fit to harbour similar relics as
this Brigg boat, while our great national store-house refuses a gift of
such importance? I make no apology to the reader for giving in detail
the result of this Brigg discovery, for it is one of the finest if not the
most instructive of any craft of this kind that has come to light in
Northern Europe. An interesting account has been written by the
Rev. D. Cary-Elwes, son of the above, and to this I am indebted for
some of the following facts.[28]

Fig. 25. The Viking Boat dug up at Brigg. Lincolnshire.

The boat is hollowed out of one huge oak log, which, from the
dimensions given above, would necessitate a tree 18 feet in
circumference, and of such a height that the branches did not begin
until 50 feet from the ground. Such a tree would be gigantic. The
bows are almost a semi-circle when viewed from above, and are
rounded off gradually to the bottom and sides, the latter being about
two inches thick and the bottom four inches. The stern, however, is
no less than sixteen inches. The transom has had to be fixed
separately on to the trunk, and the difficulty was to perform such a
piece of shipbuilding so as to make this part of the vessel as strong
and water-tight as the sides and bottom. The caulking of the joints
has been done with moss, the transom fitting into a groove across
the floor. In order that the sides of the ship might not give, in bad
weather, Mr. Cary-Elwes thinks, a tight lashing was thrown across
from one side to the other, coming round abaft of the stern, and so
keeping both sides and transom tightly together. This transom was
found a little distance away from the boat and is 4 feet wide at the
top and 2 feet 5¾ inches deep, there being a projection some 2 feet
aft, beyond the transom, so as to form an overhanging counter.
Along the whole length of the boat, close to the upper edge, holes,
2 inches in diameter, have been pierced at irregular intervals of
about 2 feet. It is uncertain what these were intended for. Although
there are no such evidences as a step for the mast, to indicate
whether she was a sailing boat, it is not safe to condemn her as
having merely been propelled by paddles. There are evidences of
decks and seats, and the primitive man would, no doubt, after he
had learned to harness the wind, maintain his mast in position
perhaps by thongs to the seat or by means of the decking. It has
even been thought that the fragment of rounded wood found with the
boat and already alluded to as a probable bowsprit, was a mast. To
me this latter supposition seems more likely than the theory of a
bowsprit. It has also been surmised that the holes running along the
boat were either for lacing to keep the ship’s sides from coming
asunder or for receptacles of pegs to hold washboards in bad
weather. Personally, I think the latter is the more probable, for it was
a very early custom. We have, in a former chapter, mentioned it as
being a practice on the Mediterranean in classical times, and we
shall see presently that the Vikings also used this method for
keeping out the spray. It happens also to be the custom among
modern savages.
Evidently during her career of activity this vessel had the
misfortune to spring a leak, for she has been patched, and the work
of the boatbuilder is most interesting to us of to-day. On the
starboard bilge a rift of 12 feet long has been made. To repair this,
wooden patches and moss have been used. The biggest patch is 5
feet 8 inches long and 6½ inches wide in the middle, tapering
gradually to a point at either end, and is of oak. The patch was let
into the rift from the outside until perfectly flush with the outer part of
the boat. On the inner side of the patch, three cleats a foot long and
four inches deep, with a hole in the centre of each, have been
attached. Wooden pins were passed through these holes, so that
pressing firmly against the solid wood on either side of the rift, they
kept the repair in position. Besides this, holes three-eighths of an
inch in diameter were made along the outer edges of the patch,
corresponding holes being also made in the fabric of the boat by
means of which the patch could be sewn to the ship with thongs.
This custom, it seems to me, would have survived in the most
natural manner from the time when the shipbuilder sewed the seams
of his skin boat. Finally, all holes and crannies were caulked with
moss. Mr. Cary-Elwes has carefully preserved a small portion of this
lacing material, which appears to be of some animal substance, and
probably twisted sinews. He has also taken some of this caulking
moss from the boat and finds that it is of two kinds, both of which
grow on sandy soils in woods, and are now largely used in the
manufacture of moss-baskets and artificial flowers.
The important fact must not be lost sight of that while all the
repairs have been made either by wood pegs or thongs, not a trace
of metal was found in the fabric of the boat. This coincides with the
argument that we have been proceeding on, viz., that such ships as
these belong not to the age of metals but to that of stone. And, as if
to convince those who scoff at the possibility of being able to fell
trees—and oak trees especially—by means of stone implements, Mr.
Cary-Elwes refers to the interesting fact that the Australian
aborigines, a type of humanity as low and primitive as one could
wish to find, had all their tools of agriculture, war and forestry, made
of stone or wood, iron being unknown to them; yet indeed they knew
how to fell the giants of the forest, such a tree as the Jarrah red gum,
now used for paving London streets, being every bit as hard as our
oak. “Within quite recent times,” adds the same author, “the
inhabitants of the South Sea Islands worked exclusively with stone
implements. I came across a good collection of these old time
weapons in New Zealand, and what is more to the point here, sundry
canoes and boats hollowed by their means. My father, who was with
me, and who is a member of the Society of Antiquaries, and not
unlearned in these matters, pointed out to me not only the similarity
that existed between these stone weapons and the prehistoric adzes
and axes of the stone age, but also the interesting fact that the
canoes hollowed out by fire or stone tools were as cleanly cut and as
cleverly wrought as the old Brigg boat.” The same writer, from the
evidence of the geological strata where she was found, concludes
that the age of the Brigg boat must be between 2600 and 3000
years, which would bring the date to between 1100 and 700 b.c.
In addition to the Brigg boat other dug-outs have been found in
various parts of our country. In 1833 one was discovered near the
river Arun in Sussex. Her length was 35 feet, breadth 4 feet, depth 2
feet. Her sides and bottom were between 4 and 5 inches thick. There
are also other similarities to the Brigg boat. In 1863 a smaller, but
similar boat, 8 feet 2 inches by 1 foot 9 inches, was also unearthed.
She had washboards like those we have attributed to the Brigg boat.
Another craft a foot smaller still was found near Dumfries in 1736,
containing a paddle. In 1822 near the Rother in Kent an immense
ship of this class measuring 63 feet long, and 5 feet broad was
unearthed also. It is interesting to remark that it was caulked with
moss in the manner already described. On the south bank of the
Clyde another of these craft was found having an upright groove in
the stern similar to that in which the sternboard of the Brigg boat was
fixed. There is also a twenty-five footer in the Museum at York.
This Brigg Boat, and the Valdermoor one, probably belong to the
class ascribed by Tacitus[29] in 70 a.d. to the Batavians and Frisians.
Some have also thought that it was in such boats as these that the
Romans crossed from Gaul to Britain. At any rate there can be no
doubt that boats of this kind were to be found at this time still existing
in Britain and along the shore washed by the English Channel and
North Sea.
In addition to those dug-outs already enumerated, a similar craft
was found in 1876 in Loch Arthur, about six miles west of Dumfries.
She is 42 feet long and like all the others is hollowed out of oak. Her
width and other characteristics show her to resemble very closely the
Brigg boat, and accentuate still more the existence of a prevailing
type of craft in Northern Europe during prehistoric times. The prow,
like that of the primitive Koryaks, is shaped after the head of an
animal. Unfortunately not the whole of this relic is preserved, but at
least one third of her, and that the bow end, is to be found in the
Museum of the Antiquarian Society of Edinburgh. More than twenty
canoes of this same class have also been found in the
neighbourhood of Glasgow. Almost all were formed out of single
tree-trunks of oak and afford evidence of having been hollowed out
by blunt tools such as the people of the Stone or Bronze Age would
possess. Two obviously later boats were dug up in 1853 and were
found to be of more elaborate construction, planks having now been
introduced. The prow resembled the beak of an ancient galley, the
stern being formed of a triangular piece of oak. For fastening the
planks to the ribs oak pins and metallic nails had been used. For
caulking, wool dipped in tar had been employed. Boehmer in his
exceedingly valuable and careful paper on “Prehistoric Naval
Architecture of the North of Europe,”[30] to which I am greatly
indebted for some important facts, points out that in the bottom of
one of these canoes a hole had been closed by means of a cork-
plug, which Professor Geikie remarks could only have come from the
latitudes of Spain, Southern France, or Italy. The inference is, of
course, that notwithstanding their island home, even the very early
inhabitants of Great Britain were in communication with distant parts
of the Continent.
There can be no doubt that, at any rate among the least
progressive peoples of Northern Europe, this dug-out, monoxylon
type of boat lasted till very late, for an account is given by Velleius
Paterculus, who about the year 5 a.d. served under Tiberius as
prefect of cavalry. He distinctly refers to the Germanic craft as dug-
outs, “cavatum, ut illis mos est, ex materia.” Pliny the elder speaks of
the piratical ships of the Chauci, one of the most progressive of the
coast tribes of Northern Europe, as having visited the rich provinces
of Gallia. These ships were dug-outs and carried thirty men. This fact
is interesting, as being the first time the Teutons had ventured on the
open sea.
During the years 1885 to 1889, while excavations were being
made at the port of Bremen at the mouth of the Weser, as many as
seven of these dug-outs were found in the alluvial land at depths of
from 6½ feet to 13 feet below the present level of the surface. They
were made of oak-trunks, and had apparently been fashioned by
axes. They were as usual flat-bottomed, without keels, but with prow
cut obliquely and with holes for the insertion of oars. Of the seven
four were entirely demolished, but of the remaining three the
dimensions were respectively: 35 feet long by 2 feet 6 inches wide;
33 feet 4 inches long by 3 feet 6 inches; 26 feet 7 inches by 3 feet 3
inches. The height varied from 1 foot 5 inches up to 2 feet 2 inches.
Several specimens of this type are preserved in the municipal
museum of Bremen.
So much, then, for the earliest type of craft. We have seen that the
dug-out in the course of time became strengthened with ribs. The
next stage in the advancement of the prehistoric shipbuilder is to
dispense with the strenuous work which necessitated the hollowing
out of a whole tree trunk of hard oak. The affixing of ribs has given
him an idea. So, utilising the hides of the wild animals which he has
shot whilst hunting, he stretches these over the same framework that
he had used for strengthening his oak-trunk. He is still in the Stone
Age, so nails are not yet invented. The skins have to be sewn
together to fit the framework, and the result is precisely that of the
coracle even now used in Wales and off Connemara. If the reader
should happen never to have seen one of these, a visit to the
Victoria and Albert Museum will quickly clear up any
misunderstanding. Though we have no actual specimens of ancient
skin-ships existing—and indeed we should not expect such a relic—
yet the interesting survival of the boat-building language of that
primitive time is found both in the Norwegian and English language
of to-day. Thus, when you have allowed a ship to lie high and dry in
the summer sun so that the planking warps and daylight can be seen
through, what is the expression you would use to express this?
Would you not remark that she has opened her seams? Now “seam”
is an Anglo-Saxon word connoting the joining together of two edges
of some texture by means of a needle. But let us take a further
instance. Do you not constantly hear shipbuilders and designers
refer to the planking that covers a ship’s ribs as her skin? Thus we
have still in common use the very words which our sires employed in
reference to the sewn hides of their primitive craft. Indeed, when one
considers that all through history, even until now, shipbuilding has
been an industry apart from ordinary occupations, and that both
ships and seamen are, as we said in our introductory chapter, the
most conservative of all peoples or created things, this survival is not
so unnatural as might seem at first. We could continue to give other
examples in the pertinacity of ancient seafaring expressions, but that
would only be to digress from the immediate subject before us. We
need only make reference to the interesting fact that Cæsar during
his first Spanish campaign in the civil war, when he required some
boats at the banks of the river Sicoris to get across, ordered the
soldiers to make boats of the build that they had learned in former
years from the British use. Thus first the keel was obtained and ribs
were fashioned of light stuff; the rest of the boat’s body being then
woven together of osiers and finally covered with hides.[31]
According to Pliny the Britanni also in the first century of our era put
to sea in wicker vessels done round with a covering of ox hide. In
such vessels they would take a six days’ voyage to the Island of
Mictis, whence the tin came.
We come now to the Bronze and Iron Ages. With the advent of
metals we find a revolution scarcely inferior to that caused by the
discovery of the edged stone. For whereas the latter could cut, yet its
efforts were confined within narrow limitations. It was capable of
felling a tree and of hollowing out its trunk with the expenditure of
considerable labour and tediousness, yet that was its highest
achievement in the department of shipbuilding. But now that the
introduction of metals, of iron and bronze, is made, the primitive man
finds that his sphere of energy is vastly widened. Instead of
hollowing out the tree he cuts it up into planks. Instead of having to
sew the outside together with thongs of hide, he has metallic nails as
fastenings. To the same kind of ribs that framed his skin-boat, he can
now nail down planks of oak and fir. He has a lighter and more easily
propelled boat than the dug-out, and a stronger and more seaworthy
ship than that made of stretched skins, although it is only fair to
observe that the hide-boat was capable of far more than one would
suppose. Mr. Jochelson in the account of the Jesup Expedition
already referred to, relates his experience of being taken for a sail in
one of the skin-boats of the Koryak. He was delighted by the
endurance which the skins (of seal) exhibited. Not the least
remarkable feature was the fact that the skin was capable of
sustaining enormous weights without bursting. But in Europe our
ancestors must have been glad to be able to discard the hide for that
of wood, since the wear and tear in beaching on rock, pebble, or
snag, exposed them to instant uselessness.
Although shipbuilding proper comes with the Metallic Age, we
must not assume that the change was made universally or at once.
The transition would be made rapidly or but slowly in proportion as
the tribe or nation were enthusiastically maritime or otherwise. In
some parts of Europe the skin-boat or even the dug-out would be in
use, while other shores were seeing built vessels of planks and ribs.
The first historic account that we possess of these more modern
vessels is to be found in Cæsar’s account of the Naval Campaign
against the Veneti in the year 54 b.c. From this narrative we learn
that the ships of the Veneti were somewhat flatter than those of the
Romans, so that they could more easily encounter the shallows and
ebbing of the tide.[32] The prows, we are told, were raised very high,
and the sterns likewise—“proræ admodum erectæ atque item
puppes”—so that they were suited for the force of the waves and
storms which they had been constructed to sustain. We have, then,
here a new design in naval architecture recorded—the Viking type of
ship—although it had been in existence for a considerable time in
the North. The high prows and sterns would immediately impress
those who had come from the more peaceful waters of Italy. Further
it is said that these ships were built of oak throughout and designed
to be enormously strong. The crossbeams, made of logs a foot thick,
were fastened by iron spikes as thick as a man’s thumb. The
anchors were made fast by iron chains instead of cables, while their
sails were made of skin and dressed leather. These were used
because they lacked canvas or the knowledge to apply it to such a
use, or more probably because they thought canvas would be of too
little strength to endure the tempests of the ocean and violent gales
of wind, and that ships of such great burden could not be managed
by them. Perhaps in the use of hides for sails, we have the parent of
the practice of using tanned sails so common in our fishing fleet and
barges. The relative character of the two kinds of ships Cæsar points
out, as we mentioned earlier in the chapter, was that the Roman fleet
excelled in speed alone and in oarsmanship. Otherwise the ships of
the Veneti were, considering the nature of the place, and the
violence of the storms more suitable and better adapted on their
side. Nor could the Roman ships injure severely the ships of the
Veneti by means of their beaks, so strong were they. And further, so
high were these ships that the Romans found great difficulty in
hurling weapons at them. Whenever a storm arose and the ships of
the Veneti ran before the wind, they could weather it more readily
and heave-to safely in the shallows, and when left by the tide feared
nothing from rocks and shelves, for—“the risk of all such things,”
ends the account pathetically, “was much to be dreaded by our
ships.”
Those who are familiar with the terrible tides and treacherous
coast of northern France[33] will readily understand how such able
Viking-like ships as the Veneti possessed, appealed to the Romans
with their fast but unsuitable craft. The difference would be that
between the smart Thames skiff and the tubby though seaworthy
dinghy of a North Sea fishing smack. For we know pretty accurately
now, thanks to the Althiburus mosaics referred to in the previous
chapter, just what Cæsar’s craft were like. Hitherto we have known
them as naves actuariæ—that is, light vessels of surpassing speed.
But if the reader will refer back to Fig. 24 he will find that the navis
actuaria, whilst propelled both with oars and sail, was nevertheless
not much of a ship to be caught in off the rocks and narrow channels
in a breeze of wind. Although these actuariæ were neither freight
ships (onerariæ) nor war-vessels properly speaking, yet they still
possessed rams and were used on this expedition for a war-like
purpose. There cannot be much doubt that the Veneti had obtained
their design and ideas of shipbuilding from the Norsemen who
relentlessly swept down from their colder climes and plundered and
pillaged from one end of the coast of Northern Europe to the other.
As we shall see presently, this design was prevalent for many years
before Cæsar came, and as we shall also see from the following
chapter it had altered but little at the time when William the
Conqueror left the French shores for England in the eleventh
century.
In the year 15 a.d. we learn from Tacitus[34] that Germanicus had
built near the mouth of the river Rhine a thousand ships with sharp
bows so as to be able to resist better the waves. Some had flat
bottoms to enable them to take the ground with impunity. Some had
a steering apparatus at both bow and stern in order that thereby they
could be rowed in either direction. Many were decked for the
accommodation of throwing machines. They were equally useful as
rowing and sailing ships, and just as in the mediæval times ships
were built with towering decks for “majesty and terror of the enemy,”
so as early as this period these vessels were imposing as to their
size whilst inspiring confidence to their own soldiery. Good
serviceable ships as they were, yet after defeating the Cherusci at
the mouth of the Ems they were shipwrecked in a storm although the
wind blew from the south. It is only fair to add, however, that the
ancients, especially the Romans, were wont to build their vessels
very quickly[35] and consequently they erred, no doubt, in
constructing them too slightly. The Saxons who, after the death of
Alexander the Great, came to the mouth of the Elbe and subjugated
the Thuringians, and who are said to have possessed the art of
tacking, already referred to, had such light vessels as belonged to
the stone age. They were wonderfully light, made out of willows and
covered with skins, but had a keel of knotty oak; yet these daring
navigators, without compass or chart, and with but a feeble
knowledge of the stars, managed to find their way to the Orkneys.
We pass now from the English Channel and the Rhine to consider
that land which has given birth to a long line of robust, vigorous ships
and men, who after the Phœnicians are the finest race of seamen
that ever sailed a sea. A little clumsy like their ships the
Scandinavians have always throughout history stood for manliness
and strength. And if we were right when we submitted that a nation’s
character exhibits itself in a most marked degree in its ships, surely
of no people could this remark be made with greater truth than
concerning the inhabitants of that Northern peninsula who, in the
early days of our own country, harassed our forefathers beyond all
endurance, but left behind to us the heritage of a love of the sea.[36]
There is in the Viking ship and its descendants not so much beauty
as nobility, not prettiness but power. The first mention of these
Northerners is by Tacitus[37] who refers to them as the Suiones.
(Tacitus died a.d. 108.) As Cæsar was struck by the difference
between the Roman ships and those of the Veneti, so Tacitus
remarks that the ships of the Suiones differ from the Romans’, too.
Although these were not sailing ships—nec velis ministrantur—yet
they were of the same design as those which were fitted with mast
and sail. Double-ended, they could easily be beached and in battle
could the more rapidly manœuvre ahead or astern.

Fig. 26. Ancient Scandinavian Rock-carving, showing Viking Ship-

forms.

But we have much earlier information than the writings of the

Roman chronicler. We have history written in stone, obvious,
illustrative and imperishable. In many parts of the Scandinavian