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VetBooks.ir

Manuel Yúfera Editor

Emerging
Issues in Fish
Larvae Research
 Emerging Issues in Fish Larvae Research
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VetBooks.ir

Gilthead seabream larva (42 days after hatching). Photo: Bernd Ueberschär
 Manuel Yúfera
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Editor

Emerging Issues in Fish

Larvae Research

123
 Editor
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Manuel Yúfera
Instituto de Ciencias Marinas de Andalucía,
(ICMAN-CSIC), Campus Universitario
Río San Pedro s/n
Puerto Real, Cádiz
Spain

ISBN 978-3-319-73243-5 ISBN 978-3-319-73244-2 (eBook)

https://doi.org/10.1007/978-3-319-73244-2
Library of Congress Control Number: 2017962104

© Springer International Publishing AG 2018

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part
of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations,
recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission
or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar
methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this
publication does not imply, even in the absence of a specific statement, that such names are exempt from
the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the
authors or the editors give a warranty, express or implied, with respect to the material contained herein or
for any errors or omissions that may have been made. The publisher remains neutral with regard to
jurisdictional claims in published maps and institutional affiliations.

Printed on acid-free paper

This Springer imprint is published by Springer Nature

The registered company is Springer International Publishing AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
 Preface
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Fish larvae are amazing organisms. They are among the smallest free-living forms
of vertebrates and exhibit fascinating growth potential. In fish aquaculture, all of the
phases of the life cycle from the fertilized egg to the adults are essential, and if one
of them fails, the whole production cycle fails. The larval phase, however, has
traditionally been considered the most sensitive period both in the wild and for
cultured fish. The vulnerability at this stage has been a powerful driving force for
fundamental and applied research. In this research line, the horizon is to know what
is necessary to allow the fish larvae to grow and develop properly and to become
healthy. The development of basic larviculture procedures based on feeding with
rotifers and Artemia in the 1970s and 1980s allowed a constant supply of fry to
support the increasing fish farming production during the last decades. Research on
fish larvae biology has continued since then and has been aimed at increasing our
basic knowledge and improving rearing methodologies. However, there are still
large knowledge gaps and the rearing process is far from optimal. For years, sci-
entific research has focused on relevant limiting factors seriously affecting survival
and growth, in a sequential manner. New species have been introduced by applying
similar protocols, and only some aspects related to feeding and nutrition have
received scientific attention. This simplistic empirical approach is insufficient for
understanding mechanisms for development and interrelations with the surrounding
rearing water. During the last decade, new analytical tools have opened up new
avenues for research in both physiology and the influence of environmental con-
ditions. In the past few years, books and specialized journals have published
reviews on the different disciplines related to fish larvae biology and aquaculture,
providing wide descriptions of main topics like development, pathologies, nutrition,
feeding, and physiology. These publications have increased our knowledge from a
textbook perspective. It may thus be considered that there is overall, good
knowledge on larval fish biology and on the current reality of larval rearing. Most
of this information, however, is based on a small number of species, and knowledge
in larval biology is advancing fast. Nowadays, there are new research ideas that
have implications not only for fish larvae but also for other vertebrates, and there
are new approaches to old problems, using omics methodologies, for example.

v
 vi Preface

The research effort on larvae is now moving beyond the strict interest for growing
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fish; it is also focusing on using them as model organisms for fundamental research.
Moreover, the numerous species being studied within the context of aquaculture
will help to uncover both the variability and the similarities within this diverse
group of vertebrates.
Instead of providing broad overviews covering general topics in larval fish
biology, this book presents examples of novel research in fish larvae with very
specific objectives. It refers to current advances derived from recent projects and a
doctoral thesis targeted at filling specific gaps in our knowledge. By representing
alternative points of view, the different chapters show how rearing and environ-
mental conditions affect physiology and developmental processes from a molecular
basis, and how these factors influence the final characteristics of late larvae and
juveniles. Overall, this book will point to recent findings on the importance of
environmental cycles, some specific nutrients, and the microbial environment on
developmental processes. There are more emerging topics of interest, but with these
few examples we hope to illustrate the dynamism of current research within
this field. These are exciting times for biologists and the discipline of biology—
especially when the target of the research is fish larvae.

Puerto Real, Spain Manuel Yúfera

 Contents
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1 Investigating Fish Larvae-Microbe Interactions in the 21st

Century: Old Questions Studied with New Tools . . . . . . . . . . . . . . 1
Ragnhild I. Vestrum, Birgit Luef, Torunn Forberg, Ingrid Bakke
and Olav Vadstein
2 Environmental Cycles and Biological Rhythms During Early
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Francisco Javier Sánchez-Vázquez and José Fernando López-Olmeda
3 The Digestive Function in Developing Fish Larvae and Fry. From
Molecular Gene Expression to Enzymatic Activity . . . . . . . . . . . . . 51
Manuel Yúfera, Francisco J. Moyano
and Gonzalo Martínez-Rodríguez
4 Variability in Digestive Enzyme Capacity in Early Stages of
Marine Fish Larvae: Ontogenetic Variations, Biorhythms,
Hormonal Control and Nutrient Sensing Mechanisms . . . . . . . . . . 87
Bernd Ueberschär, Carmen Navarro-Guillén, Ana Gomes,
Ivar Rønnestad, Carlos Rojas-Garcia, Inken Hanke,
Dagh Sommerfeld and Robert Tillner
5 Phospholipids in Marine Larval Rearing . . . . . . . . . . . . . . . . . . . . 131
Keshuai Li, Rolf Erik Olsen, Yang Jin and Yngvar Olsen
6 Fat-Soluble Vitamins in Fish: A Transcriptional Tissue-Specific
Crosstalk that Remains to be Unveiled and Characterized . . . . . . . 159
Ignacio Fernández, Paulo Gavaia, Maria J. Darias and Enric Gisbert
7 Nutritional Modulation of Marine Fish Larvae Performance . . . . . 209
Sofia Engrola, Cláudia Aragão, Luisa M. P. Valente
and Luís E. C. Conceição

vii
 viii Contents

8 Fish Pigmentation. A Key Issue for the Sustainable Development

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of Fish Farming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229

Laura Cal, Paula Suarez-Bregua, Paloma Moran,
José Miguel Cerdá-Reverter and Josep Rotllant
9 Novel Aspects of Phosphate Endocrine Control: A Key Element
for the Long-Term Sustainability of Finfish Aquaculture . . . . . . . . 253
Paula Suarez-Bregua, Laura Cal, Pedro M. Guerreiro
and Josep Rotllant
10 Feeding and Development of Warm Water Marine Fish Larvae in
Early Life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Jing Hu, Yibing Liu, Zhenhua Ma and Jian G. Qin
 Chapter 1
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Investigating Fish Larvae-Microbe

Interactions in the 21st Century:
Old Questions Studied with New Tools

Ragnhild I. Vestrum, Birgit Luef, Torunn Forberg, Ingrid Bakke

and Olav Vadstein

Abstract All animals need a mutualistic interaction with their microbiota for
proper development and functioning. Also for the fish-microbiota interaction con-
siderable research has been done, and especially for reared fish larvae this inter-
action is crucial for their viability. However, during the 1980s and 1990s a number
of findings revealed at that time current methods were not suitable for studying the
total microbial community and that data on composition of microbiota was biased.
Several recent methodological revolutions have boosted the possibilities for
addressing questions related to fish larvae-microbiota interactions that previously
lacked suitable tools for proper evaluation. These methodological achievements
include the development of experimental rearing systems including gnotobiotic
systems for fish, new visualization tools, and molecular “omics” tools for charac-
terizing the response of the host on a variety of levels and for characterizing both
composition and activity of fish microbiota. We present and review these tools and
give examples on how they have been used to improve our understanding of fish
larvae-microbiota interactions. With respect to understanding, this includes in
particular how the microbiota is established and maintained, what the functionality
of the microbiota is and how it affects fish health, and finally how we can apply this
knowledge for management of a healthy and beneficial microbiota in aquaculture
settings.

Keywords Microbiome Germ-free model systems
Imaging


Molecular methods Omics

R. I. Vestrum
B. Luef
T. Forberg
I. Bakke
O. Vadstein (&)

Department of Biotechnology and Food Science, NTNU Norwegian
University of Science and Technology, 7491 Trondheim, Norway
e-mail: [email protected]
Present Address:
T. Forberg
BioMar AS, Pirsenteret, 7010 Trondheim, Norway

© Springer International Publishing AG 2018 1

M. Yúfera (ed.), Emerging Issues in Fish Larvae Research,
https://doi.org/10.1007/978-3-319-73244-2_1
 2 R. I. Vestrum et al.

1.1 Introduction
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Mariculture has the possibility of being a major contributor to feeding a growing

human population, but this will require solutions to significant challenges (Duarte
et al. 2009). A major biological challenge will be to overcome bottlenecks with
regards to producing high quality juveniles suitable for the intensive cultivation
environment. We know that e.g. egg quality, nutrition and physiochemical condi-
tions contribute to the observed problems, but the well-known observation of poor
reproducibility between fully replicated tanks with equal genetic variability (e.g.
full sibling groups) cannot be explained by such factors (Vadstein et al. 1993). Even
though this is a complex problem with multiple dimensions, it is now demonstrated
beyond any doubt, that detrimental host-microbe interactions are a major cause of
these problems (Vadstein et al. 1993, 2013).
Since Robert Kock and Louis Pasteur’s major discoveries in the 1880s, the
relationship between man and their microbes has had primary focus on disease.
However, during the last 15 years a paradigm shift has occurred, as substantial
effort has been devoted to better understanding the role of our microbiota for health
and normal development. The last decennium has revealed a multitude of ways in
which the microbiota affects the normal development of animals, including fish
(Kanther and Rawls 2010; Sekirov et al. 2010). Most of this research has been
motivated by human health and therefore most studies have been done in man and
mammalian model organisms. It is now important to verify the generality of these
findings with other vertebrates and invertebrates, including fish as they have a core
role in the evolution of vertebrates.
Until 15 years ago quantitative and qualitative studies of microbes in aquacul-
ture systems were performed by culture-based techniques, and until the 60s it was a
general consensus that the knowledge of microbes associated with animals was
fairly well understood. Culture-dependent techniques mean that samples are serially
diluted and spread on so-called non-selective and selective medium on agar plates.
Quantitative estimates of densities are then given as colony forming units (CFU).
Since the late 1970s it has been shown that the discrepancy between total counts in
epifluorescence microscope and CFU is huge—typically a factor of 1000 for natural
waters. This is the first problem for culture-dependent methods. For some time it
was claimed that this difference was due to dead microbes, but it is now well
established that this can only account for a limited part of the discrepancy (e.g.
Karner and Fuhrman 1997). For man-made systems, like aquaculture systems, the
culturability can be considerably higher than in natural systems, but the difference
in the estimated abundance of microbes between the two methods is normally still
one order of magnitude. Studies during the last two decades have shown that the
difference between CFU and total counts is due to the selectivity of non-selective
agar—i.e. the majority of the microbes are still not possible to culture at laboratory
conditions (Hugenholtz et al. 1998). This is known as the “great plate anomaly”
(Hugenholtz 2002) and points to a second problem with culture-based methods; for
qualitative analysis of the composition of the microbes, the culture-based results are
 1 Investigating Fish Larvae-Microbe Interactions … 3

highly biased, as the culturability is dependent on phylogeny (Hugenholtz et al.

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1998). A third drawback of culture-based methods is that it is time consuming. Pure

cultures have to be established from colonies through several steps of sub-culturing,
and finally the pure cultures must be identified through a substantial number of
tests. For appropriate identification of species composition more than 100 pure
cultures should be characterized per sample (1% resolution = 1 isolate represents
1% of the population). In reality, few studies have characterized that many pure
cultures per sample, which is a clear indication of the labor involved. For one of the
authors of this chapter (O.V.) these methodological limitations felt overwhelming in
the early 90s. However, since then the revolution in method development has
caused another type of paradigm shift. This shift is largely due to the rapid
development in molecular biology and imaging methods. Method development,
possibilities, and applications are covered in detail below.
Based on the knowledge of the bias in culture-based studies one question is
unpleasant but impossible to disregard: Do we have to do it all over again? Or more
specific, is the cultivation-based knowledge we have on host-microbe interactions
in fish so biased that it counteracts the progress in our knowledge and under-
standing? The answers to these questions are not straight forward. First, all types of
“old” knowledge is probably not equally biased. Second, even though results from
high-throughput sequencing studies of fish microbiota are accumulating at a decent
speed, the amount of data is still too limited to do a proper evaluation of “old” data.
The severity of the problem is exemplified by the study of Fjellheim et al. (2012)
who found a negative correlation between quantification of bacteria associated with
larvae by plating on agar plates versus by quantitative PCR of rDNA. Thus we are
convinced that we have to do a lot of the work over again.
The aim of this chapter is to give an overview of new methods and how they can
be used to speed up and improve our understanding of fish-microbiota interactions.
With respect to understanding, this includes in particular how the microbiota is
established and maintained, what the functionality of the microbiota is and how it
affects fish health, and finally how we can apply this knowledge for management of
a healthy and beneficial microbiota in aquaculture settings. We will describe the
methods with focus on applications without going into technical details, and give
examples of applications for studying host-microbiota interactions.

1.2 Experimental Designs for Studying Fish-Microbiota

Interactions

Traditionally, experiments with fish are run in cultivation tanks. These are normally
just a downscaling of aquaculture production systems, and therefore realistic sys-
tems from an aquaculture perspective. However, the degree of experimental control
for fish-microbe studies is limited. Below we will describe different experimental
systems, going from high relevance/low control to low relevance/high control.
 4 R. I. Vestrum et al.

We want to stress that it is important to keep this range of experimental systems, as

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high control systems can be used to study mechanisms and hypotheses which then
can be verified in systems where the fish live a more normal life, i.e. low control
systems. We will try to highlight pros and cons within the different systems.

1.2.1 Traditional Rearing Tanks

Traditional rearing tanks can have variable volumes and can be run in traditional
flow-through systems (FTS) or as recirculating aquaculture systems (RAS), where
the water is reused after removal of waste and toxic substances. Tank volumes
depend on the size of the fish and the population size needed for experimental
reasons. From an ethical point of view and due to legislation the number of animals
used in an experiment should be as low as possible, but especially in applied
research the experimental systems should be relevant for a practical setting. This
might create a conflict between ethics and the practical relevance of new knowl-
edge. The use of FTS versus RAS has clear implications for fish-microbe studies, as
for RAS there will also be a recirculation of microbes. In some situations this can be
an advantage (maintaining the effect of the fish on the microbiota), whereas in other
cases a disadvantage (a high background concentration of microbes). FTS versus
RAS is therefore an important part of the experimental design. Traditional rearing
tanks have limited possibilities to control the microbiota in the rearing water. This
includes import and export of microbes, but particularly microbial growth in the
rearing tanks. For example, in studies with probiotics, it might be difficult to predict
the probiotic microbes’ ability to establish in the system as the competitive situation
with background microbes will vary depending on the species present and system
design. However, we have shown that it is possible to control and stabilize the
microbiota in rearing tanks through a selection regime against some microbes, and
favoring others (r- and K-selection) (Attramadal et al. 2014).

1.2.2 Systems with Rearing of Single Individuals

One of the drawbacks of using normal rearing tanks is that you have limited
possibilities for controlling import and export of microbes. Another problem is that
one fish may affect the other individuals in the same tank. This is important as
moribund individuals may infect healthy individuals, and that there is a continuous
sharing of microbes by defecation and re-ingestion (Reitan et al. 1998). A way out
of this problem is to rear single individuals in small units. Size and type of units
may vary from e.g. 40 ml in plastic cups (Forberg et al. 2016) to 2 ml in
24-multiwell plates (Fjellheim et al. 2010; Sandlund and Bergh 2008). Size of the
system may vary dependent on species and the length of the experiment. These
types of systems can be run without water exchange for yolk sac stages and early
 1 Investigating Fish Larvae-Microbe Interactions … 5
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Fig. 1.1 Survival of yolk-sac larvae of Atlantic cod (Gadus morhua) incubated together with
different bacteria, including background bacteria (positive control), a pathogen (negative control)
and three probiotics candidates, versus time. Each treatment included 72 individuals. Data from
Fjellheim et al. (2010)

first feeding (Fjellheim et al. 2010; Sandlund and Bergh 2008) or with an irregular
water exchange for longer experiments (Forberg et al. 2016). In systems with water
exchange, there are possibilities for maintaining some degree of control of the
microbes entering the system by e.g. adding a defined or described microbiota. It is,
however, important to keep in mind that the way by which the renewal of the water
takes place may create different types of selection regimes for the microbiota in the
rearing unit (e.g. continuous versus periodic dilution rate).
An example of the use of 24-multiwell plates with single individuals in 2 ml of
water is shown in Fig. 1.1. In this experiment, unfed yolk-sac larvae were exposed
to either probiotic candidates, a pathogen (negative control) or only background
bacteria (positive control). As shown in the figure, you get reliable results with a
limited number of individuals (in this case 72 individuals per treatment), and the
reproducibility is very good (unpublished results). This simple system easily
revealed that one probiotic candidate actually was pathogenic (ID 4-29). In systems
with rearing of single individuals, it is possible to reduce the number of individuals
used for the experiment to a minimum, and confidence intervals for the survival can
be calculated from surviving individuals and the total number of individuals based
on binomial statistics (Box et al. 1978). Moreover, chi-square and exact tests can be
used for statistical analysis.
The pros of these systems should be evident based on the information above.
The down side is mainly the simplicity of the system and for some species the
difficulty of rearing the fish for sufficiently long periods. The simplicity includes
lack of interactions between individuals of fish, which is an advantage for some
 6 R. I. Vestrum et al.

types of experiments. This clearly emphasizes the advantage of having a possibility

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for doing experiments in different types of systems, and selecting the systems based
on the aim of the study.

1.2.3 Germ-Free and Gnotobiotic Systems

If traditional rearing tanks is on one end of the experimental system scale, germ-free
and gnotobiotic systems are on the other end. A germ-free system has no microbes
at all, and in a gnotobiotic system the biota in the system is known (= gnotos) and
predefined. This can only be achieved by first generating germ-free test animals or
fish, which are then deliberately colonized by one or several bacterial strains known
to the researcher. Gnotobiotic systems were first developed for mammals such as
rats, but also for fish several gnotobiotic systems have been published (Table 1.1).
In general, the procedure of creating germ-free fish involves a first step with surface
disinfection of the eggs and subsequent hatching in an environment without
microbes. The subsequent rearing under germ-free or gnotobiotic conditions may be
in the presence or absence of antibiotics. When rearing the fish with antibiotics, you
can only use antibiotic resistant bacteria and there is also a possibility of negative
effects on larvae due to long term exposure to antibiotics (Moullan et al. 2015). As
documented in Table 1.1, many different techniques have been used to surface
disinfect the eggs, and gnotobiotic systems have been attempted for several fish
species.
The main advantage of using these systems is the high degree of control of the
microbial environment. This is appealing for many types of experiments with a
reductionistic approach—a strategy with a long history of success in natural sci-
ence. The main disadvantage of germ-free and gnotobiotic systems is the com-
plexity in performing these experiments. A reproducible method for achieving
germ-free larvae is needed, and avoiding contamination during the experiment is
also crucial (i.e. maintain them germ-free or gnotobiotic). In the cases were the fish
is fed live prey, the prey also needs to be germ-free. When using Artemia as live
feed it is possible to hatch them from disinfected cysts, and this makes the process
more straightforward. However, when using e.g. rotifers a germ-free culture has to
be established, they have to be reared germ-free, and therefore must be fed
germ-free feed (microalgae or yeast). Moreover, this production line has to be kept
throughout the germ-free or gnotobiotic fish experiment. One problem not given
much attention in the literature, is the maintenance of the gnotobiotic condition.
When only one microbe is added it is a presence/absence problem, but when several
species are added in a given ratio it is not straight forward to maintain this ratio.
This has to do with the selection regimes in the rearing unit and due to renewal of
water (see also Sect. 1.2).
Results generated from one fish species may not be easily transferrable to
another, as germ-free zebrafish (Danio rerio) seem to be less developed than
conventional fish (Rawls et al. 2004), while germ-free sea bass larvae have more
Other documents randomly have
different content
 The town is utterly gone. There are those who Hippo
argue that this or that was not done as history
relates, because of this or that no vestige remains; and if tradition
tells them that Rome built here or there, they deny it, because they
cannot find walls, however much they dig (within the funds their
patrons allow them). These men are common in the universities of
Europe. They are paid to be common. They should see Hippo.
Here was a great town of the Empire. It detained the host of
Vandals, slaves and nomads for a year. It was the seat of the most
famous bishopric of its day, and within its walls, while the siege still
endured, St. Augustine died. It counted more than Palermo or
Genoa: almost as much as Narbonne. It has completely
disappeared. There are not a few bricks scattered, nor a line of
Roman tiles built into a wall. There is nothing. A farmer in his
ploughing once disturbed a few fragments of mosaic, but that is all:
they can make a better show at Bignor in the Sussex weald, where
an unlucky company officer shivered out his time of service with
perhaps a hundred men.
In the heart of the Tell, behind the mountains Calama
which hide the sea, yet right in the storms of the
sea, in its clouds and weather, stands a little town which was called
Calama in the Roman time and is now, since the Arabs, called
Guelma.
It is the centre of that belt of hills. A broad valley, one of the
hundreds which build up the complicated pattern of the
Mediterranean slope, lies before the platform upon which the fortress
rose. A muddy river nourishes it, and all the plain is covered with the
new farms and vineyards—beyond them the summits and the
shoulders that make a tumbled landscape everywhere along the
northern shores of Africa guard the place whichever way one turns.
From the end of every street one sees a mountain.
If a man had but one day in which to judge the nature of the
province, he could not do better than come to this town upon some
winter evening when it was already dark, and wake next morning to
see the hurrying sky and large grey hills lifting up into that sky all
around and catching the riot of its clouds. It is high and cold: there is
a spread of pasture in its fields and a sense of Europe in the air. No
device in the architecture indicates an excessive heat in summer and
even the trees are those of Italy or of Provence. Its site is a survival
from the good time when the Empire packed this soil with the cities
of which so great a number have disappeared: it is also a promise of
what the near future may produce, a new harvest of settled and
wealthy walls, for it is in the refounding of such municipalities that the
tradition of Europe will work upon Africa and not in barren adventure
southward towards a sky which is unendurable to our race and under
which we can never build and can hardly govern.
 Guelma is typical in every way. It was Berber before the Romans
came, but nothing remains of its founders or of whatever punic
influence its first centuries may have felt. Of Rome so much endures
that the heavy walls and the arches are, as it were, the framework of
the place.
In the citadel a great fragment larger than The Permanence of
anything else in the town runs right across the Rome
soldiers’ quarters, pierced with the solid arches that once supported
the palace of Calama. Only the woodwork has disappeared. The
stones which supported the flooring still stand out unbroken, and the
whole wall, though it is not very high—hardly higher than the big
barracks around it—remains in the mind, as though it had a right to
occupy one’s memory of the Kasbah by a sort of majesty which
nothing that has been built since its time has inherited. Here, as
throughout the Empire, the impression of Rome is as indefinable as
it is profound, but one can connect some part of it at least with the
magnitude of the stones and the ponderous simplicity of their
courses, with the strength that the half-circle and the straight line
convey, and with the double evidence of extreme antiquity and
extreme endurance; for there is something awful in the sight of so
many centuries visibly stamped upon the stone, and able to evoke
every effect of age but not to compel decay.
This nameless character which is the mark of the Empire, and
carries, as it were, a hint of resurrection in it, is as strong in what has
fallen as in what stands. A few bricks built at random into a mud wall
bear the sign of Rome and proclaim her title: a little bronze
unearthed at random in the rubbish heaps of the Rummel is a
Roman Victory: a few flag-stones lying broken upon a deserted path
in the woodlands is a Roman Road: nor do any of these fragments
suggest the passing of an irrecoverable good, but rather its
continued victory. To see so many witnesses small and great is not
to remember a past or a lost excellence, but to become part of it and
to be conscious of Rome all about one to-day. It is a surety also for
the future to see such things.

There is a field where this perpetuity and this The Peasant’s Wall
escape from Time refresh the traveller with
peculiar power. It is a field of grass in the uplands across which the
wind blows with vigour towards distant hills. Here a peasant of the
place (no one knows when, but long ago) fenced in his land with
Roman stones. The decay of Islam had left him aimless, like all his
peers. He could not build or design. He could not cut stone or mould
brick. When he was compelled to enclose his pasture, the only
material he could use was the work of the old masters who had
trained his fathers but whom he had utterly forgotten or remembered
only in the vague name of “Roum.” It was long before the reconquest
that he laboriously raised that wall. Some shadow of Turkish power
still ruled him from Constantine. No one yet had crossed the sea
from Europe to make good mortar or to saw in the quarries again. It
is with a lively appreciation that one notes how all he did is perishing
or has perished. The poor binding he put in has crumbled. The slabs
slope here and there. But the edges of those stones, which are
twenty times older than his effort, remain. They will fall again and lie
where he found them; but they and the power that cut them are alike
imperishable.
It has been said that the men of antiquity had The Landscape of
no regard for landscape, and that those principal Antiquity
poems upon which all letters repose betray an indifference to
horizons and to distant views. The objection is ill-found, for even the
poems let show through their admirable restraint the same passion
which we feel for hills, and especially for the hills of home: they
speak also of land-falls and of returning exiles, and an Homeric man
desired, as he journeyed, to see far off the smoke rising from his
own fields and after that to die. But much stronger than anything
their careful verse can give us of this appetite for locality is the
emplacement of their buildings.
Mr. March-Phillips has very well described the The Theatre of
spirit which built a certain temple into the Calama
scenery of a Sicilian valley. Here (he says), in a place now deserted,
the white pillars ornament a jutting tongue of land, and are so placed
that all the lines of the gorge lead up to them, and that the shrine
becomes the centre of a picture, and, as it were, of a composition. Of
this antique consciousness of terrestrial beauty all southern Europe
is full, and here in Guelma, upon an edge of the high town, the site of
the theatre gives evidence of the same zeal.
 The side of a hill was chosen, just where the platform of the city
breaks down sharply upon the plain below. There, so that the people
and the slaves upon the steps could have a worthy background for
their plays, the half-circle of the auditorium was cut out like a quarry
from the ground. Beyond the actors, and giving a solemnity to the
half-religious concourse of the spectators, the mountains of the Tell
stood always up behind the scene, and the height, not only of those
summits but of the steps above the plain, enhanced the words that
were presented. We have to-day in Europe no such aids to the
senses. We have no such alliance of the air and the clouds with our
drama, nor even with our patriotism—such as the modern world has
made it. The last centuries of the Empire had all these things in
common: great verse inherited from an older time, good statuary,
plentiful fountains, one religion, and the open sky. Therefore its
memory has outlasted all intervening time, and it itself the Empire,
(though this truth is as yet but half-received,) has re-arisen.

There is one great note in the story of our race The Greatness of
which the least learned man can at once The First Four
appreciate, if he travels with keen eyes looking Centuries
everywhere for antiquity, but which the most learned in their books
perpetually ignore, and ignore more and more densely as research
develops. That note is the magnitude of the first four centuries.
Everybody knows that the ancient world ran down into the
completed Roman Empire as into a reservoir, and everybody knows
that the modern world has flowed outwards from that reservoir by
various channels. Everybody knows that this formation of a United
Europe was hardly completed in the first century, that it was at last
conscious of disintegration in the fifth. The first four centuries are
therefore present as dates in everybody’s mind, yet the significance
of the dates is forgotten.
Historians have fallen into a barren contemplation of the Roman
decline, and their readers with difficulty escape that attitude. Save in
some few novels, no writer has attempted to stand in the shoes of
the time and to see it as must have seen it the barber of Marcus
Aurelius or the stud-groom of Sidonius’ Palace. We know what was
coming, the men of the time knew it no more than we can know the
future. We take at its own self-estimate that violent self-criticism
which accompanies vitality, and we are content to see in these 400
years a process of mere decay.
The picture thus impressed upon us is certainly false. There is
hardly a town whose physical history we can trace, that did not
expand, especially towards the close of that time. There was hardly
an industry or a class (notably the officials) that had not by an
accumulation of experience grown to create upon a larger and a
larger scale its peculiar contribution to the State; and far the larger
part of the stuff of our own lives was created, or was preserved, by
that period of unity.
That our European rivers are embanked and canalised, that we
alone have roads, that we alone build well and permanently, that we
alone in our art can almost attain reality, that we alone can judge all
that we do by ideas, and that therefore we alone are not afraid of
change and can develop from within—in a word, that we alone are
Christians we derive from that time.
Our theory of political justice was partly formulated, partly handed
on, by those generations; our whole scheme of law, our conceptions
of human dignity and of right. Even in the details our structure of
society descends from that source: we govern, or attempt to govern,
by representation because the monastic institutions of the end of the
Empire were under a necessity of adopting that device: we associate
the horse with arms and with nobility because the last of the Romans
did so.
If a man will stand back in the time of the Antonines and will look
around him and forward toward our own day, the consequence of the
first four centuries will at once appear. He will see the unceasing
expansion of the paved imperial ways. He will conceive those great
Councils of the Church which would meet indifferently in centres
1500 miles apart, in the extremity of Spain or on the Bosphorus: a
sort of moving city whose vast travel was not even noticed nor called
a feat. He will be appalled by the vigour of the western mind between
Augustus and Julian when he finds that it could comprehend and
influence and treat as one vast State what is even now, after so
many centuries of painful reconstruction, a mosaic of separate
provinces. He will calculate with what rapidity and uniformity the
orders of those emperors who seem to us the lessening despots of a
failing state were given upon the banks of the Euphrates, to be
obeyed upon the Clyde. He will then appreciate why the Rome which
Europe remembers, and upon which it is still founded, was not the
Rome of literature with its tiny forum and its narrow village streets,
but something gigantic like that vision which Du Bellay had of a
figure with one foot upon the sunrise and its hands overspreading
ocean.
Indeed this great poet expresses the thing more vividly by the
sound of three lines of his than even the most vivid history could do.

“Telle que dans son char la Bérycynthienne

Couronnée de tours, et heureuse d’avoir
Enfanté tant de dieux....”

This was the might and the permanence from which we sprang.
To establish the character of the Empire and its creative mission is
the less easy from the prejudice that has so long existed against the
action of religion, and especially of that religion which the Empire
embraced as its cataclysm approached. The acceptation of the
creed is associated in every mind with the eclipse of knowledge and
with a contempt for the delights which every mind now seeks. It is
often thought the cause, always the companion, of decay, and so far
has this sentiment proceeded that in reading books upon Augustine
or upon Athanasius one might forget by what a sea and under what
a sunlight the vast revolution was effected.
 It is true that when every European element had mixed to form
one pattern, things local and well done disappeared. The vague
overwhelming and perhaps insoluble problems which concern not a
city but the whole world, the discovery of human doom and of the
nature and destiny of the soul, these occupied such minds as would
in an earlier time have bent themselves to simpler and more feasible
tasks than the search after finality. It is true that plastic art, and to a
less extent letters, failed: for these fringes of life whose perfection
depends upon detail demand for their occasional flowering small and
happy States full of fixed dogmas and of certain usages. But though
it lost the visible powers antiquity had known, the Empire at its end,
when it turned to the contemplation of eternity, broadened much
more than our moderns—who are enemies of its religious theory—
will admit. The business which Rome undertook in her decline was
so noble and upon so great a scale that when it had succeeded,
then, in spite of other invasions, the continuity of Europe was saved.
We absorbed the few barbarians of the fifth century, we had even the
vitality to hold out in the terror and darkness of the ninth, and in the
twelfth we re-arose. It was the character of the Western Empire
during the first four centuries, and notably its character towards their
close, which prevented the sleep of the Dark Ages from being a
death. These first four centuries cast the mould which still constrains
us; they formed our final creed, they fixed the routes of commerce
and the sites of cities, and perpetually in the smallest trifles of
topography you come across them still: the boundary of Normandy,
as we know it to-day, was fixed by Diocletian. If there can be said of
Europe what cannot be said of any other part of the world, that its
civilisation never grew sterile and never disappeared, then we owe
the power of saying such a thing to that long evening of the
Mediterranean.

If this pre-eminence of Rome in the process of The Arabic

her conversion is the lesson of all travel it is Influence
especially the lesson of Africa; and nowhere is that lesson taught
more clear than in Guelma. Here also you may perceive how it was
that the particular cause which ruined the spirit of the Roman town
also saved its stones, and you may feel, like an atmosphere, the
lightness, the permeation, as it were, without pressure:—the
perpetual fluid influence which overflowed the province upon the
arrival of the Arabs. So that the bone of Rome remain, caught in a
drift of ideas which, like fine desert sand, could preserve them for
ever.
For the Arab did in Calama what he did throughout Barbary: he
cast a spell. He did not destroy with savagery, he rather neglected all
that he could afford to neglect. Here also he cut down timber, but he
did not replant. Here also he let the water-pipes of the Romans run
dry. Here also the Arab, who apparently achieved nothing material,
imposed a command more powerful than the compulsion of any
government or the fear of any conqueror: he sowed broadcast his
religion and his language; his harvest grew at once; first it hid and at
last it stifled the religion and the language he had found. The
speech, and the faith which renders that speech sacred, transformed
the soul of Barbary: they oppose between them a barrier to the
reconquest more formidable by far than were the steppes and the
nomads to the first advance of Rome. Of this impalpable veil which is
spread between the native population and the new settlers the
traveller is more readily aware in the little cities of the hills than in the
larger towns of the coast. The external change of the last generation
is apparent: the houses about him are European houses; the roads
might be roads in France or Northern Italy. The general aspect of
Guelma confirms that impression of modernity, nor is there much
save the low loop-holed walls which surround the town, to remind
one of Africa; but from the midst of its roofs rises the evidence of that
religion which still holds and will continue to hold all its people. The
only building upon which the efforts of an indolent creed have
fastened is the mosque, and the minaret stands alone, conspicuous
and central over all the European attempt, and mocks us.
Far off, where the walls and the barracks are confused into a
general band of white, and no outline is salient enough to distinguish
the modern from the ancient work of the place, this wholly
Mohammedan shaft of stone marks the place for Mohammedan. It is
an enduring challenge.
 There is a triumph of influence which all of us have known and
against which many of us have struggled. It is certainly not a force
which one can resist, still less is it effected by (though it often
accompanies) the success of armies. It is the pressure and at last
the conquest of ideas when they have this three-fold power: first, that
they are novel and attack those parts of the mind still sensitive;
secondly, that they are expounded with conviction (conviction
necessary to the conveyance of doctrine); and, thirdly, that they form
a system and are final. Such was the triumph of the Arab.
Our jaded day, which must for ever be taking some drug or tickling
itself with unaccustomed emotion, has pretended to discover in
Islam, as it has pretended to discover in twenty other alien things,
the plan of happiness; and a stupid northern admiration for whatever
has excited the wonder or the curiosity of the traveller has made
Mohammedism, as it has made Buddhism and God knows what
other inferiorities or aberrations of human philosophy, the talk of
drawing-rooms and the satisfaction of lethargic men. It is not in this
spirit that a worthy tribute can be paid to the enormous invasion of
the seventh century.
That invasion as a whole has failed. The Arabic
Christendom, for ever criticised, (for it is in its Invasion
own nature to criticise itself,) has emerged; but if one would
comprehend how sharp was the issue, one should read again all that
was written between Charlemagne and the death of St. Louis. In the
Song of Roland, in the “Gesta Francorum,” in Joinville, this new
attack of Asia is present—formidable, and greater than ourselves;
something which we hardly dared to conquer, which we thought we
could not conquer, which the greatest of us thought he had failed in
conquering. Islam was far more learned than we were, it was better
equipped in arms and nevertheless more civic and more tolerant.
When the last efforts of the crusades dragged back to Europe an evil
memory of defeat, there was perhaps no doubt in those who
despaired, still less in those who secretly delighted that such
fantasies were ended—there was no doubt, I say, in their minds that
the full re-establishment of our civilisation was impossible, and that
the two rivals were destined to stand for ever one against the other:
the invader checked and the invaded prudent; for, throughout the
struggle we had always looked upon our rivals at least as equals and
usually as superiors.
It is in the most subtle expressions that the Its Continued
quarrel between the two philosophies appears. Influence
Continually Islam presses upon us without our knowing it. It made
the Albigenses, it is raising here and there throughout European
literature at this moment notes of determinism, just as that other
influence from the Further East is raising notes of cruelty or of
despair.
There is one point in which the contact The Gothic
between these master-enemies and ourselves is
best apparent. They gave us the Gothic, and yet under our hands
the Gothic became the most essentially European of all European
things. Consider these two tiers of one Arabian building founded in
Africa, while yet the vigour of that civilisation was strong. True, the
work is not in stone but in plaster, for to work stone they needed an
older civilisation than their own. But see how it is the origin of, or
rather identical with, our ogive. By what is it that we recognise these
intersecting segments (which are of the perfect 60° like our own) to
be something foreign? And how is it that we know that no Christian
could have built these things? Venice has windows like these: by just
so much she is not of the West, and by just that innoculation perhaps
she perished. The ecstasy of height, the self-development of form
into further form, the grotesque, the sublime and the enthusiastic—
all these things the Arab arch lacks as utterly as did the Arab spirit;
yet the form is theirs and we obtained it from them. In this similarity
and in these differences are contained and presented visibly the
whole story of our contact with them and of our antagonism.
 In the presence of the doom or message which the Arabians
communicated to our race in Africa, one is compelled to something
of the awe with which one would regard a tomb from which great
miracles proceeded, or a dead hero who, though dead, might not be
disturbed. The thing we have to combat, or which we refrain and
dread from combating, is not tangible, and is the more difficult to
remove. It has sunk into the Atlas and into the desert, it has filled the
mind of every man from the Soudan which it controls up northwards
to Atlas and throughout this land.
Roaming in the Sahara are bands of men The Touaregs
famous for their courage and their isolation.
They are called the Touaregs. They are of the same race and the
same language as those original Berbers who yet maintain
themselves apart in the heights of Aurès or of the Djurdjura. They
are the enemies of all outside their tribes, especially of the Arab
merchants, upon whose caravans they live by pillage. Yet even these
Islam has thoroughly possessed and would seem to have conquered
for ever. Their language has escaped; their tiny literature (for they
have letters of their own, and their alphabet is indigenous) has
survived every external influence, but even there the God of the
Mohammedans has appeared.
 One taken captive some years since wrote back from Europe to
his tribe in his own stiff characters a very charming letter in which he
ended by recommending himself to the young women of his home,
for he himself was a fighter, courteous, and in his thirtieth year. But
when he had written “Salute the Little Queens from me,” he was
careful to add an invocation to Allah. And if in their long forays it is
necessary to bury hastily some companion who has fallen in the
retreat, his shallow grave in the sand is carefully designed according
to the custom of religion. They leave him upon his right side in an
attitude which they hold as sacred, his face turned to the east and
towards Mecca. In this posture he awaits the Great Day.
Against this vast permanent and rooted The Lack of an
influence we have nothing to offer. Our designs Opposing Faith
of material benefit or of positive enlightenment are to the presence of
this common creed as is some human machine to the sea. We can
pass through it, but we cannot occupy it. It spreads out before our
advance, it closes up behind. Nor will our work be accomplished until
we have recovered, perhaps through disasters suffered in our
European homes, the full tradition of our philosophy and a faith
which shall permeate all our actions as completely as does this faith
of theirs.
That no religion brought by us stands active against their own is
an apparent weakness in the reconquest, but that consequence of
the long indifference through which Europe has passed is not the
only impediment it has produced. The dissolution of the principal
bond between Europeans—the bond of their traditional ritual and
confession—has also prevented the occupation of Africa from being,
as it should have been, a united and therefore an orderly campaign
of the West to recover its own.
Had not our religion suffered the violent Cause of Isolated
schisms which are now so slowly healing, and French Action
had not our general life resolved itself for a time into a blind race
between the various provinces of Europe, the reconquest of Barbary
would have fallen naturally to the nations which regard each its own
section of the opposing coast; as in the reconquest of Spain the
Asturias advanced upon Leon, the Galicians upon Portugal, and Old
Castille upon the southern province to which it extended its own
name. Then Italy would have concerned itself with Tunis—with
Ifrigya, that is—and with the rare fringe of the Tripolitan and its
shallow harbours. The French would have occupied Numidia. The
Spaniards would have swept on to re-Christianise the last province
of the west from Oran to the Atlantic, and so have completed the
task which they let drop after the march upon Granada. Such should
have been the natural end of mediæval progress, and that
reconstruction of the Empire (which was the nebulous but constant
goal towards which the Middle Ages moved) would have been
accomplished. But the most sudden and the most inexplicable of our
revolutions came in and broke the scheme. The Middle Ages died
without a warning. A curious passion for metaphysics seized upon
certain districts of the north, which in their exaltation attempted to
live alone: the south, in resisting the disruption of Europe, exhausted
its energies; and meanwhile the temptation to exploit the Americas
and the Indies drained the Mediterranean of adventurers and of
navies. Islam in its lethargy acquired new vigour from its latest
converts, and the Turks, with none but the Venetians to oppose
them, tore away from us the whole of the Levant and rode up the
Danube to insult the centre of the continent. The European system
flew apart, and its various units moved along separate paths with
various careers of hesitation or of fever. It was not until the
Revolution and the reconstitution of sane government among us that
the common scheme of the west could reappear.
On this account—on account of the vast disturbance which
accompanied the Reformation and the Renaissance—Europe halted
for three centuries. When at last a force landed upon the southern
shore of the Mediterranean, it was a force which happened to be
despatched by the French.
The vices and the energy of this people are The French
well known. They are perpetually critical of their
own authorities, and perpetually lamenting the decline of their
honour. There is no difficulty they will not surmount. They have
crossed all deserts and have perfected every art. Their victories in
the field would seem legendary were they not attested; their
audacity, whether in civil war or in foreign adventure, has
permanently astonished their neighbours to the south, the east and
the north. They are the most general in framing a policy and the
most actual in pursuing it. Their incredible achievements have
always the appearance of accidents. They are tenacious of the
memory of defeats rather than of victories. They change more
rapidly and with less reverence than any other men the external
expression of their tireless effort, yet, more than any other men, they
preserve—in spite of themselves—an original and unchanging spirit.
Their boundaries are continually the same. They are acute and vivid
in matters of reason, careless in those of judgment. A coward and a
statesman are equally rare among them, yet their achievements are
the result of prudence and their history is marked by a succession of
silent and calculating politicians. Alone of European peoples the
Gauls have, by a sort of habit, indulged in huge raids which seemed
but an expense of military passion to no purpose. They alone could
have poured out in that tide of the third century before our era to
swamp Lombardy, to wreck Delphi, and to colonise Asia. They alone
could have conceived the crusades: they alone the revolutionary
wars. It is remarkable that in all such eruptions they alone fought
eastward, marching from camp into the early light; they alone were
content to return with little spoil and with no addition of provinces, to
write some epic of their wars.
It is evident that such a people would produce in Africa, not a
European and a general, but a Gallic and a particular effect. They
boast themselves in everything the continuators of the Romans.
They do, indeed, inherit the Roman passion for equality, and they,
like the Romans, have tenaciously fought their way to equality by an
effort spread over many hundred years. They are Roman in their
careful building, in their strict roads, in their small stature, in their
heavy chests, in their clarity of language, in their adoration of office
and of symbol, in their lightning marches: the heavy lading of their
troops, their special pedantry, their disgust at vagueness, their
ambition and their honour are Roman. But they are not Roman in
permanent stability of detail. The Romans spread an odour of
religion round the smallest functions of the State: of the French you
can say no more than that any French thing you see to-day may be
gone to-morrow, and that only France remains. They are not Roman
in the determination never to retreat, nor are they Roman in the
worship of silence. The French can express the majesty of the
Empire in art: they cannot act it in their daily life—for this inheritance
of Rome the Spaniards are better suited. As for the Roman
conception of a fatal expansion the Russians exceed them, and for
the Roman ease and aptitude the Italians.
Had, then, the reconquest of Barbary fallen naturally to the three
sisters—to Spain, to Italy, and to France, the long attempt of Europe
might have reached its end. The Spaniard would have crushed and
dominated in Morocco where the Mohammedan was most strongly
entrenched; the Italian, with his subtle admixture, would have
kneaded Tunis and the eastern march into a firm barrier; the French
would have developed their active commerce upon the many small
towns of the Central Tell, would have pierced, as they are fitted to
pierce, the high Central Plateaux with admirable roads, and would
have garrisoned, as their taste for a risk well fits them to garrison,
the outposts of the Central Atlas against the desert. Then the task
would be over, and Europe would be resettled within its original
boundaries.

On their long route marches, on the marches The March

of their manœuvres and their wars, the French,
along their roads which are direct and august, (and at evening, when
one is weary, sombre,) seek a place of reunion and of repose: upon
this the corps converges, and there at last a man may lie a long night
under shelter and content to sleep: a town lies before the pioneers
and is their goal. It stands, tiny with spires, above the horizon of their
hedgeless plains, and as they go they sing of the halt, or, for long
spaces, are silent, bent trudging under the pack: for they abhor
parade. Very often they do not reach their goal. They then lie out in
bivouac under the sky and light very many fires, five to a company or
more, and sleep out unsatisfied. Such a strain and such an attempt:
such a march, such a disappointment, and such a goal are the
symbols of their history; for they are perpetually seeking, under
arms, a Europe that shall endure. In this search they must continue
here in Africa, as they continue in their own country, that march of
theirs which sees the city ever before it and yet cannot come near to
salute the guard at the gates and to enter in. It is their business to re-
create the Empire in this province of Africa. It may be that here also
they will come to no completion; but if they fail, Europe will fail with
them, and it will be a sign that our tradition has ended.
They have done the Latin thing. First they The French Genius
have designed, then organised, then built, then
ploughed, and their wealth has come last. The mind is present to
excess in the stamp they have laid upon Africa. Their utter regularity
and the sense of will envelop the whole province; and their genius,
inflexible and yet alert, alert and yet monotonous, is to be seen
everywhere in similar roads, similar bridges of careful and even
ornamented stone, similar barracks and loop-holed walls.
There is a perspective upon the High Plateaux The Straight
which though it is exceptional is typical of their Railway
spirit. It is on the salt plain just before the gate of the desert is
reached and the fall on to the desert begun. Here the flat and
unfruitful level glares white and red: it is of little use to men or none.
Some few adventurers, like their peers in the Rockies, have
attempted to enclose a patch or two of ground, but the whole
landscape is parched and dead. Through this, right on like a gesture
of command, like the dart of a spear, goes the rail, urging towards
the Sahara, as though the Sahara were not a boundary but a goal.
The odd, single hills, as high as the Wrekin or higher, upon which not
even the goats can live, look down upon the straight line thus traced:
these hills and the track beneath them afford a stupendous contrast.
Nowhere is the determination of man more defiant against the sullen
refusal of the earth.
 There is another effort of the French which The French
may be watched with more anxiety and more Afforestation
comprehension by northern men than their admirable roads or their
railways or their wires above the sand, and that is their afforestation.
It is a debate which will not be decided (for the material of full
decision is lacking) whether, since the Romans crowded their
millions into this Africa, the rainfall has or has not changed. It is
certain that they husbanded water upon every side and built great
barricades to hold the streams; yet it is certain, also, that their cities
stood where no such great groups of men could live to-day. There
are those who believe that under Atlas, towards the desert, a shallow
sea spread westward from the Mediterranean and from Syrtis: there
are others who believe that the dry water-courses of the Sahara
were recently alive with streams, and that the tombs and inscriptions
of the waste places, now half buried in the sand, prove a great lake
upon whose shores a whole province could cultivate and live. Both
hypotheses are doubtful for this reason—that no good legend
preserves the record. Changes far less momentous have left whole
cycles of ballads and stories behind them. The Sahara has been the
Sahara since men have sung or spoken of it. Moreover, the Romans
did certainly push out, as the French have done, towards certain
limits, beyond which no effort was worth the while of armies. They
felt a boundary to the south. They could bear the summer of Biskra,