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Hermann Ehrlich Editor

Extreme
Biomimetics
Extreme Biomimetics
Hermann Ehrlich
Editor

Extreme Biomimetics

123
Editor
Hermann Ehrlich
Institute of Experimental Physics
TU Bergakademie Freiberg
Freiberg, Sachsen
Germany

ISBN 978-3-319-45338-5 ISBN 978-3-319-45340-8 (eBook)

DOI 10.1007/978-3-319-45340-8
Library of Congress Control Number: 2016950236

© Springer International Publishing AG 2017

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Dedicated to Joanna Aizenberg
Preface

Extreme Biomimetics is a novel interdisciplinary scientific direction of modern

biomimetics proposed for the first time in 2010; and is currently a vibrant area of
research. It was born at the crossroads between such scientific disciplines as
Prebiotic Chemistry and Mineralogy, Astrobiology, Evolutionary Biology,
Hydrothermal Chemistry and Biochemistry, and Exobiology. These in turn include
scientific directions such as the Primordial Soup Theory, the Origin and Evolution
of Life, and Extreme Biomineralization (Fig. 1). To delve into these fields, radical
rethinking was necessary, one that explores unusual and very unique chemical and
biochemical scenarios. Such studies could lead to both a better understanding of the
origin of life under harsh environmental conditions, and to new approaches for
designing new composite materials with entirely novel physicochemical properties.
Today, Extreme Biomimetics is well on track as a powerful approach, and a
milestone for modern biological materials inspired chemistry. This is particularly
true where there is strong interest in the combination of various inorganic
nano-organized structures with biological macromolecules; as well as using such
macromolecules as templates and scaffolds. In contrast to traditional aspects of
biomimetic synthesis of these hybrid materials at ambient temperatures, Extreme
Biomimetics is based on mineralization and metallization of various biomolecules
under conditions mimicking extreme aquatic niches like hydrothermal vents,
geothermal pipelines, or hot springs with temperatures near boiling point. Here,
organisms known as thermophiles, acidophiles, alkaliphiles, and polyextremophiles
represent the sources for bioinspiration. Thus, the basic principle of this concept is
to exploit biopolymers that are chemically and thermally stable under these very
specific conditions in vitro.
Extreme Biomimetics also includes both chemical and biochemical reactions at
very low temperatures—under so-called psychrophilic conditions—both in vivo
and in vitro. In this case Extreme Biomimetics and Cryobiology, as well as
Exobiology, are intimately intertwined. Processes which occur in deep-sea organ-
isms defined as barophiles and piezophiles fall under the umbrella of Extreme
Biomimetics.

vii
viii Preface

Fig. 1 Schematic view on the place of Extreme Biomimetics on the multidisciplinary crossroads

Thus, the goal of Extreme Biomimetics is to bring together broad variety of

extreme (from the biological point of view) chemical reactions with templates of
biological origin, and to develop the next generation of hybrid composites with
novel properties.
At present, unfortunately, there is no monograph that deals with the area of
Extreme Biomimetics and could, if required, serve as an accompaniment to a course
of lectures on the subject. I sincerely hope that this book would be of interest for
undergraduate and graduate students studying materials science, chemistry, bioor-
ganic chemistry, biochemistry, solid-state physics, biophysics, bioengineering, and
Preface ix

researchers, engineers, chemists, biologists, physicists, material scientists, and

physicians. Therefore, the targeted audience of proposed book is broad variety of
students and materials scientists associated with biocomposites, materials for
medicine, biomaterials and hydrothermal technology.
This issue on Extreme Biomimetics illustrates how structures and functions of
organisms that dwell under extreme conditions have inspired new forms of artificial
biomineralization and technology. The book includes 135 figures, 11 tables and
more than 1000 references. Authors from such countries as Germany, France,
Japan, Poland, Russia, Ukraine, and the United States contribute to 10 chapters.
The collection of chapters begins with a review by Ehrlich and Nikolaev on
psychrophiles as source for bioinspiration in biomineralization and biological
materials science. They focus attention on biosilicification (in diatoms, silicoflag-
ellates, radiolarians, sponges), and biocalcification (in bacteria, foraminiferans,
sponges, bryozoans, corals, molluscs, echinoderms, crustaceans), and pay special
attention to the biology and adaptation mechanisms of ice fish species.
Additionally, these authors consolidate a wealth of references related to the topic
that may be a time-saving resource for experts in materials science who are looking
for model or key organisms as sources for special scientific and technological
inspiration.
In continuation of this theme, Tabachnick, Janussen, and Menshenina look at a
remarkable cold biosilicification process in psychrophilic glass sponges. In this
chapter, the psychrophilic problem is discussed as it pertains to different aspects
of the life cycle of hexactinellid sponges. New data on the vertical distribution of
Hexactinellida is provided, which supports ideas put forth from previous investi-
gations; as well as new interpretations of their mortal process. A new type of
deep-sea reef construction of hexactinellid sponge Sarostegia oculata is described
for the first time.
The next chapter prepared by Nikolaev and co-authors is logically dedicated to
the phenomenon of psychrophilic calcification using artificial biomineralization
methods. The first efforts to develop calcium phosphate-based composites on
organic templates using a dual membrane diffusion method at the freezing point of
water have been reported. Galkin and Sagalevich (Chap. 4) have contributed to an
overview of hydrothermal vent fauna that represents a unique source for scientists
who are involved in investigations of ecology, zoology, and biochemistry of
extremophiles. Numerous unique underwater images represented in this chapter
should help give a better understanding of the life near hydrothermal vents.
Comparative in situ microscopic observation of cellulose and chitin in
hydrothermal conditions is described by Deguchi in Chap. 5. The experimental
results reported here have direct ramifications for developing hydrothermal biomass
conversion, hydrothermal synthesis of inorganic–organic hybrid composites, as
well as fossilization of soft-bodied organisms. Boury (Chap. 6) takes inspiration
from hydrothermal vent organisms and describes the integration of different
biopolymers and a metal oxide under selective hydrothermal conditions, with the
aim of developing new composites. This chapter emphasizes the important
x Preface

parameters of the synthesis, the impact of the biopolymers on characteristics of the

metal oxide, and in improving its performance in technological applications.
Stawski in Chap. 7 paid special attention to the thermostability of chitin because
of its potential application in hydrothermal synthesis, with the aim of creating novel
nanostructured metal oxide-based composite materials. The author reported that
chitin is still stable in the between 100 and 400 °C. Petrenko and co-authors (Chap.
8) propose a new term “bioelectrometallurgy,” which they define as the electro-
plating of biological matrices. For the first time, cell-free chitinous sponge skele-
tons, which possess a 3D network structure, were used for copper plating under
laboratory conditions. In Chap. 9, the unique structural, mechanical, and thermal
properties of chitin from the “biomaterials” point of view have been discussed by
Wysokowski, Kaiser, and Jesionowski. These authors also described the basic
principles of solvothermal synthesis and utilization of chitin as a structural template
in hydrothermal reactions. Finally, Szatkowski and Jesionowski (Chap. 10) ana-
lyzed the wide range of applications of the thermostable structural protein spongin
of Poriferan origin in terms of its utilization in Extreme Biomimetics.
It may be sufficient to say that without enthusiasm and cooperation of all authors
who have contributed to this first book on Extreme Biomimetics, this work could
hardly have been attempted. I am very grateful to the German Research Foundation
(DFG, Project EH 394/3-2) as well as to Krüger Research School,
Biohydrometallurgical Center for Strategic Elements (BHMZ) at TU Bergakademie
Freiberg, Germany, for financial support. I also thank Profs. Catherine Skinner,
Edmund Bäeuerlein, Victor Smetacek, Dan Morse, Peter Fratzl, Matthias Epple,
George Mayer, Christine Ortiz, Marcus Buehler, Andrew Knoll, Hartmut Worch,
and Dirk-Carl Meyer for their support and permanent interest in our research on
Extreme Biomimetics. To Dr. Allison L. Stelling, Dr. Mikhail V. Tsurkan and
Sarah D. Smith-Tsurkan, I am thankful for taking excellent care of manuscripts and
proofs. To my parents, my wife, and my children, I am under deep obligation for
their patience and support during the years of my scientific activity.

Freiberg, Germany Hermann Ehrlich

Contents

1 Psychrophiles as Sources for Bioinspiration

in Biomineralization and Biological Materials Science . . . . . . . . . . . 1
Hermann Ehrlich and Anton Nikolaev
2 Cold Biosilicification in Metazoan: Psychrophilic
Glass Sponges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Konstantin Tabachnick, Dorte Janussen and Larisa Menshenina
3 Psychrophilic Calcification In Vitro . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Anton Nikolaev, Vasilii V. Bazhenov, Olga V. Frank-Kamenetskaya
and Olga V. Petrova
4 Endemism and Biodiversity of Hydrothermal Vent Fauna . . . . . . . 97
Sergey V. Galkin and Anatoly M. Sagalevich
5 Comparative In Situ Microscopic Observation
of Cellulose and Chitin in Hydrothermal Conditions . . . . . . . . . . . . 119
Shigeru Deguchi
6 Biopolymers for Biomimetic Processing of Metal Oxides . . . . . . . . . 135
Bruno Boury
7 Thermogravimetric Analysis of Sponge Chitins
in Thermooxidative Conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
Dawid Stawski
8 Bioelectrometallurgy of Copper on Chitin . . . . . . . . . . . . . . . . . . . . . 205
Iaroslav Petrenko, Vasilii V. Bazhenov, Allison L. Stelling
and Valentina Z. Kutsova
9 Hydrothermal Synthesis of Advanced Chitin-Based Materials . . . . . 223
Marcin Wysokowski, Sabine Kaiser and Teofil Jesionowski

xi
xii Contents

10 Hydrothermal Synthesis of Spongin-Based Materials . . . . . . . . . . . . 251

Tomasz Szatkowski and Teofil Jesionowski
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Contributors

Vasilii V. Bazhenov Institute of Experimental Physics, TU Bergakademie

Freiberg, Freiberg, Germany
Bruno Boury ICG-CMOS UMR 5253, Université de Montpellier, Montpellier
Cedex 05, France
Shigeru Deguchi Research and Development Center for Marine Biosciences,
Japan Agency for Marine-Earth Science and Technology (JAMSTEC), Yokosuka,
Japan
Hermann Ehrlich Institute of Experimental Physics, TU Bergakademie Freiberg,
Freiberg, Germany
Olga V. Frank-Kamenetskaya Institute of Earth Sciences, Department of
Crystalography, St. Petersburg State University, St. Petersburg, Russia
Sergey V. Galkin P.P. Shirshov Institute of Oceanology of Academy of Sciences
of Russia, Moscow, Russia
Dorte Janussen Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am
Main, Germany
Teofil Jesionowski Faculty of Chemical Technology, Institute of Chemical
Technology and Engineering, Poznan University of Technology, Poznan, Poland
Sabine Kaiser Institute of Experimental Physics, TU Bergakademie Freiberg,
Freiberg, Germany
Valentina Z. Kutsova Department of Material Science the Name U.N.
Taran-Zhovnir, National Metallurgical Academy of Ukraine, Dnipropetrovsk,
Ukraine
Larisa Menshenina Physical Department, Moscow State University, Moscow,
Russia

xiii
xiv Contributors

Anton Nikolaev Institute of Earth Sciences, Department of Crystallography,

St. Petersburg State University, St. Petersburg, Russia
Iaroslav Petrenko Institute of Experimental Physics, TU Bergakademie Freiberg,
Freiberg, Germany
Olga V. Petrova Laboratory of Theoretical Physics, Mathematic Department,
Komi Science Center of the Ural Division of the Russian Academy of Science,
Syktyvkar, Russia
Anatoly M. Sagalevich P.P. Shirshov Institute of Oceanology of Academy of
Sciences of Russia, Moscow, Russia
Dawid Stawski Department of Commodity and Material Sciences and Textile
Metrology, Lodz University of Technology, Lódź, Poland
Allison L. Stelling Department of Biochemistry, Duke University, Durham, USA
Tomasz Szatkowski Faculty of Chemical Technology, Institute of Chemical
Technology and Engineering, Poznan University of Technology, Poznan, Poland
Konstantin Tabachnick P.P. Shirshov Institute of Oceanology of Academy of
Sciences of Russia, Moscow, Russia
Marcin Wysokowski Faculty of Chemical Technology, Institute of Chemical
Technology and Engineering, Poznan University of Technology, Poznan, Poland
Chapter 1
Psychrophiles as Sources
for Bioinspiration in Biomineralization
and Biological Materials Science

Hermann Ehrlich and Anton Nikolaev

1.1 Biodiversity of Psychrophilic Organisms

The Arctic and Antarctic areas are the best localities to search for psychrophilic pro-
and eukaryotes. Traditionally, antarctic fauna have been investigated from the view
of evolutionary history (Ortmann 1901; LaBrecque and Barker 1981; Watling and
Thurston 1989; Clarke and Crame 1989; Clarke 1996; Clarke et al. 2004; Rogers
2007; Whittle et al. 2014), zoogeography (Murphy 1928; Lindsey 1940a, b;
Ostapoff 1966; Hedgepeth 1969; Voss 1988; Barnes and De Grave 2001; Griffiths
et al. 2009; Brandt et al. 2014; Griffiths and Waller 2016), ecology (Littlepage and
Pearse 1962; Holdgate 1967; Headland 1989; Knox 1970; White 1984; Convey
2001; Arntz and Clarke 2002; Clarke 2003; Bowden 2005; Clarke et al. 2007;
Ducklow et al. 2007; McClintock et al. 2008, 2010; Avila et al. 2009; Gutt et al.
2015; Clark et al. 2015), life history strategies (Eastman 1993), physiology, bio-
chemistry, molecular biology, genomics, survival and adaptation mechanisms
(Pearse and Giese 1966a; Dayton et al. 1974; Richardson and Hedgpeth 1977; Brey
et al. 1996; Clark et al. 2004; Peck et al. 2005; Bowden et al. 2006; Rogers 2007;
Roterman et al. 2016), community dynamics (Blake and Narayanaswamy 2004),
productivity and reproduction (Tressler 1964; Pearse et al. 1991; Aronson et al.
2009), and biodiversity (Ehrenberg 1844a, b; Hooker 1846; Pfeffer 1889; DeWitt

H. Ehrlich (&)
Institute of Experimental Physics, TU Bergakademie Freiberg,
Leipziger str. 23, 09599 Freiberg, Germany
e-mail: hermann.ehrlich@physik.tu-freiberg.de
A. Nikolaev
Institute of Earth Sciences, Department of Crystallography,
St. Petersburg State University, Universitetskaya nab., 7-9,
199034 St. Petersburg, Russia
e-mail: floijan@gmail.com

© Springer International Publishing AG 2017 1

H. Ehrlich (ed.), Extreme Biomimetics,
DOI 10.1007/978-3-319-45340-8_1
2 H. Ehrlich and A. Nikolaev

1965; Swithinbank et al. 1961; Holme 1964; Andriashev 1965; Arnaud et al. 1967;
Bullivant 1967; Dell 1972; Gallardo 1987; Dayton 1990; Arntz et al. 1997; Gutt
and Starmans 1998; Starmans et al. 1999; Clarke and Johnston 2003; Eastman
2005; Barnes et al. 2006; Brandt et al. 2007a, b; Griffiths 2010; Costello et al. 2010;
Waller 2013; Gutt et al. 2013; Brandt et al. 2014; WoRMS Editorial Board 2015;
Griffiths and Waller 2016). However, antarctic marine bentos also represent the
amazing source for inspiration for such scientific disciplines and directions as
bioinspired materials chemistry, materials science, biomineralogy, bionic, and
biomimetics. The total number of both vertebrates and benthic invertebrate species
described and observed from the Antarctic continental shelf range between 4000
and 17,000 species (see for review Arntz et al. 1997; Ragua-Gil et al. 2004; Gutt
et al. 2004). Many of these species are endemic (Cailleux 1961), for example,
100 % of nematodes (Andrassy 1998), 90 % of amphipods, 66 % of isopods, over
90 % of pycnogonids, 73 % of echinoderms, 58 % of bryozoans, and 95 % of fish
(Dayton et al. 1974). To obtain more information about biodiversity (Fig. 1.1) and
recently discovered species, we recommend to study following databases: EASIZ
(Ecology of the Antarctic Sea Ice Zone) supported by SCAR (the Scientific
Committee on Antarctic Research), ANDEEP (Antarctic Benthic Deep-Sea
Biodiversity) as an integral project of CeDAMar (Census of the Diversity of
Abyssal Marine Life), CAML (Census of Antarctic Marine Life) and the
SCAR EBA (Evolution and Biodiversity in Antarctic), Census of Antarctic Marine
Life (CAML), and the Scientific Committee on Antarctic Research Marine
Biodiversity Information Network (SCAR-MarBIN scientific programs (Gili et al.
2001; Brandt et al. 2004; Brandt and Hilbig 2004; Griffiths 2010; Kennicutt et al.
2014).

Fig. 1.1 The biodiversity of psychrophilic biomineralyzers on the bottom of selected areas of
Antarctic seas is amazing. Reprinted from Gutt (2004a), photograph: J. Gutt ©AWI/MARUM
(Germany), published under: Creative Commons Attribution 3.0 Unported
1 Psychrophiles as Sources for Bioinspiration … 3

1.2 Psychrophilic Biosilicification

The new term “psychrophilic biosilicification” was first proposed in 2015 (Ehrlich
2015). The research subjects in this case are psychrophilic silica-producing uni-
cellular (silicoflagellates, diatoms) and multicellular organisms like sponges (both
Demospongiae and Hexactinellida). According to Maldonado and co-workers, our
current understanding of the silicon cycle in the ocean assumes that diatoms
dominate not only the uptake of silicic acid, but also the production and recycling of
biogenic silica (Maldonado et al. 2005). Other organisms with siliceous skeletons,
including sponges, radiolarians, and silicoflagellates, are thought to play a negli-
gible role. The authors showed that the retention of Si by siliceous sponges in some
sublittoral and bathyal environments is substantial, and that sponge populations
function as Si sinks. Therefore, sponges may affect Si cycling dynamics and Si
availability for diatoms, particularly in Si-depleted environments. It was strongly
suggested (Maldonado et al. 2005) that the role of sponges in the benthopelagic
coupling of the Si cycle is significant. For example, Antarctic giant hexactinellids,
such as Rossella nuda (Fig. 1.2) and Scolymastra joubini (Fig. 1.3), which may be
up to 2 m tall, 1.4 m in diameter, and up to 600 kg wet weight, can contain up to
50 kg biogenic silica each (Maldonado et al. 2005). These sponges represent
excellent examples of scientific phenomenon known as “polar gigantism” (Chapelle
and Peck 1999). Psychrophilic biosilicification definitively occurs also in both
Arctic and Antarctic species of silicoflagellates, radiolarians, ebridians (Korhola
and Smol 2001), and diatoms. The precipitation of silica by all of these organisms
depresses dissolved silica concentrations in shallow marine waters (Nelson and
Smith 1986; Nelson et al. 1995; DeMaster 2002; Maldonado et al. 2011; Tréguer
and De La Rocha 2013).

Fig. 1.2 Antarctic glass sponge Rossella nuda. Reprinted from Gutt (2004b), photograph: J. Gutt
©AWI/MARUM (Germany), published under: Creative Commons Attribution 3.0 Unported
4 H. Ehrlich and A. Nikolaev

Fig. 1.3 Antarctic glass sponge Scolymastra joubini. Reprinted from Gutt and Teixidó (2004),
photograph: J. Gutt ©AWI/MARUM (Germany), published under: Creative Commons Attribution
3.0 Unported

1.3 Psychrophilic Diatoms

Materials scientists are fascinated by diatoms especially due to ability of these

unicellular microalgae to construct nano- and microscaled, highly perforated
siliceous skeletons with a broad diversity of fascinating architectures. The cell walls
(frustules) of photosynthetic (Kuczynska et al. 2015) diatoms are both mechanically
stable (Hamm et al. 2003) and transparent to light. Principles of biosilicification in
diatoms have been under intensive investigations over the last 20 years. The pos-
sible role of numerous biosilica-associated organic matrices based on peptides and
special proteins (Kröger and Sumper 1998; Sumper and Kröger 2004; Kröger and
Poulsen 2007; Kotzsch et al. 2016) as well as on polysaccharides (Brunner et al.
2009; Ehrlich and Witkowski 2015) is well discussed in the literature.
Unfortunately, biosilicifications in diatoms have not been studied under extreme
low (diatoms habituating within ice) nor extreme high (thermotolerant diatoms in
hot springs) temperatures. Because both light and temperature effects on metabo-
lism and bioactivity of these algae are crucial (Fiala and Oriol 1990; Ingebrigtsen
et al. 2016), the future research should be oriented on understanding of mechanisms
of biosilica formation in diatoms living in extreme conditions.
Thus, psychrophilic diatoms have been reported to live in Arctic (Grunow 1884;
Cleve 1898; Foged 1953, 1981; Meguro et al. 1966; Van Baalen and O’Donnell
1983; Abelmann 1992a; von Quillfeldt 2000; Arrigo et al. 2012; Pla-Rabés et al.
2016) and Antarctic (Bunt 1964; Mock and Junge 2007; Lyon and Mock 2014)
areas, for example, on ice crystals, ice-melt ponds (Mock and Hoch 2005), in a
brine solution in microfissures between ice crystals on the underside of the ice
1 Psychrophiles as Sources for Bioinspiration … 5

Fig. 1.4 Freshwater psychrophilic diatom Aulacoseira baicalensis

(Meguro et al. 1966), and in the interstitial water of the ice matrix formed of the
bottom surface of thick ice (Burkholder and Mandelli 1965) of fresh water
(Babanazarova et al. 1996) (Fig. 1.4) and sea water origin (Garrison and Buck
1985; Baldi et al. 2011). Some diatoms possess cold-adapted photosynthesis (Mock
and Valentin 2004) and can be localized in the pack ice, where they could reach
such high concentrations that their photosynthetic pigments discolor the ice
yellow-brown, or brown (Thomas and Dieckmann 2002). The psychrophilic diatom
Chaetoceros neogracile is known as major biomass producer that can thrive in
extreme cold environments (Hwang et al. 2008). It was shown experimentally
(Mock and Hoch 2005) that such polar diatoms as Fragilariopsis cylindrus could
efficiently acclimate photosynthesis over a wide range of polar temperatures given
enough time. Although the rate of biosilicification in Antarctic diatoms remains
unknown, some species, like Fragilariopsis kerguelensis, show high rate of
biosilicification and contain thick-shelled frustules. Interestingly, these diatoms
from the iron limited Antarctic Circumpolar Current sequester silicon, but com-
paratively little carbon (Assmy et al. 2013).
Similar to other psychrophilic organisms, diatoms produce exopolymeric sub-
stances to protect themselves against dehydration stress and the presence of ice
crystals (for review see Palmisano et al. 1985; Raymond et al. 1994; Raymond
2000; Raymond and Knight 2003; Krembs et al. 2002). For example, non-marine
psychrophilic diatoms Berkeleya and Navicula ice-active substances can inhibit the
recrystallization of ice at concentrations of approximately 1 µg/ml (total protein
plus carbohydrate) (Raymond et al. 1994). From other side, it was shown that
6 H. Ehrlich and A. Nikolaev

exudates separated from cells of the marine diatom Thalassiosira pseudonana

nucleate ice (Wilson et al. 2015).
Also, diverse ice-binding (Janech et al. 2006; Krell et al. 2008) and antifreeze
proteins (AFP), which decrease the freezing point noncolligatively, referred to as
thermal hysteresis (TH) activity, and inhibit ice recrystallization, have been recently
reported in diatoms like C. neogracile (Gwak et al. 2010) and Fragilariopsis
cylindrus (Bayer-Giraldi et al. 2011; Uhlig et al. 2011). The finding of AFP genes
(afps) as a multigene family in the diatoms genus Fragilariopsis (Bayer-Giraldi
et al. 2010) indicates the importance of this group of genes for cryosurvival of these
microalgae. Unfortunately, the possible role of AFP in biosilicification of psy-
chrophilic diatoms is still uninvestigated.

1.4 Psychrophilic Silicoflagellates

Silicoflagellates are enigmatic unicellular planktonic marine organisms with both

photosynthetic and heterotrophic activities which are not as abundant as diatoms.
They produce not very complex internal silica-based skeletons in the form of a
network of bars (see for details Preisig 1994; van Tol et al. 2012; McCartney et al.
2014), and use so-called undulipodium (flagellum) to propel themselves in the
aquatic environment. It seems that some species of silicoflagellates are highly
specialized for cold water niches. For example, Distephanus medianoctisol domi-
nates over 71 % of total silicoflagellates in the central Arctic (Takahashi et al.
2009). Numerous Arctic (Bukry 1981, 1985; Okazaki et al. 2003; Tsutsui and
Takahashi 2009) as well as Antarctic (Mandra 1969; Haq and Riley 1976;
McCartney and Wise 1988; Abelmann and Gersonde 1991) silicoflagellates species
have been described from ecological and systematics point of view. However,
publications on biosilicification phenomenon in this psychrophilic species remain
unknown. Due to possible growth of some silicoflagellates in culture (Van
Valkenburg and Norris 1970), corresponding experiments with selected species
under psychrophilic laboratory conditions seem to be of interest with respect to
isolation and investigation of skeletal structures which can be produced by the
organisms.

1.5 Psychrophilic Radiolarians

In contrast to silicoflagellates, some groups of radiolarians (i.e., Acantharea) pro-

duce unicellular mineralized skeleton made not only of silica but of strontium
sulfate (celestite) (Hollande and Martoja 1974; Nigrini and Moore 1979). However,
diverse species of Nassellaria and Spumellaria (subphylum Polycistina) both
produce ornate siliceous skeletons (tests) (for review see Anderson 1981;
Afanasieva and Amon 2014). Remarkably, in these groups the development of
1 Psychrophiles as Sources for Bioinspiration … 7

skeleton is initiated outside the central capsule. It is suggested (Harper and Knoll
1975) that during evolution natural selection, mediated by the role of the diatoms in
the global silica cycle, apparently favors radiolarian morphology which uses less
silica in test construction.
Psychrophilic radiolarians which habituate in Antarctic area are known since
first reports from nineteenth century published by Ehrenberg and Haeckel (see for
review Ehrenberg 1844a, b; Haeckel 1887). Abundance, ecology, and taxonomy of
these psychrophilic protozoans are well described (Popofsky 1908; Riedel 1958;
Hays 1965; Petrushevskaya 1967, 1975, 1986; Goll and Björklund 1974; Chen
1975; Boltovskoy and Riedel 1979; Morley and Stepien 1985; Tibbs and Tibbs
1986; Gowing 1989; Lazarus 1990; Abelmann 1992b; Nishimura et al. 1997;
Lazarus et al. 2005). One of the typical Antarctic taxa of radiolarians is named
Antarctissa spp. (Petrushevskaya 1986). Radiolarians are also to be reported in
Arctic seas (Molina-Cruz and Bernal-Ramirez 1996; Bjørklund et al. 1998;
Bjørklund and Kruglikova 2003). Although some attempts to understand biomin-
eralization in radiolarians have been carried out (Enriques 1931; Cachon and
Cachon 1971; Afanasieva 2006; Suzuki et al. 2013; Ogane et al. 2009, 2010, 2014),
there is still a lack of knowledge about the underpinning mechanisms of biosili-
cification in these protists. Probably, cultivation techniques (see for details
Matsuoka 1992) will give us the opportunity to obtain enough amounts of radio-
larian biomass under low temperature conditions for bioanalytical in-depth
investigations.

1.6 Psychrophilic Sponges

Skeletal structures of numerous psychrophilic demosponges (Figs. 1.5 and 1.6) as

well as all glass sponges are made of hierarchically organized biocomposites
(Fig. 1.7) which contain silicon dioxide and some kind of proteinaceous (silicateins,
glassin, collagen), or polysaccharide-based (chitin) organic matrix. Especially
hexactinellids have been reported as sources for bioinspiration in diverse papers in
structural biology (Barthel 1995; Leys et al. 2007; Ehrlich 2011; Shimizu et al.
2015), materials science (Aizenberg et al. 2005; Weaver et al. 2007; Ehrlich et al.
2010a, b, 2011), and optics (Cattaneo-Vietti et al. 1996; Sundar et al. 2003; Müller
et al. 2006; Ehrlich et al. 2016). However, the mechanism of psychrophilic silici-
fication is still unknown. On the basis of this gap in knowledge, combined with
renewed interest in phenomenon of biosilicification in nature, an improved
understanding of silica cycling in psychrophilic marine environments is critical.
Sponges together with diatoms represent examples of biogenic silica that have
been synthetized under psychrophilic conditions in Antarctic waters (Sañé et al.
2013). There are reports on diverse Antarctic parasitic diatoms which are able to
complete their entire biological cycle inside the sponge body (McClintock 1987;
Gaino et al. 1994; Kunzmann 1996; Hamilton et al. 1997; Bavestrello et al. 2000;
Cerrano et al. 2000, 2004). Bavestrello et al. (2000) suggested that this unusual
8 H. Ehrlich and A. Nikolaev

Fig. 1.5 Antarctic demosponges (a, b) commonly contain no pigments. Reprinted from Gutt
(2004a), photograph: J. Gutt ©AWI/MARUM (Germany), published under: Creative Commons
Attribution 3.0 Unported

symbiotic behavior could be a special strategy that allows the diatom to survive in
darkness. It seems that psychrophilic sponges have not lost their ability to syn-
thesize a broad variety of substances which they actively use for chemical defense
under low temperatures against parasites like diatoms, and even spongivorous
animals (see for review Seldes et al. 1990a, b; Baker et al. 1995; Yang et al. 1995;
Amsler et al. 2000; Moon et al. 2000; Furrow et al. 2003; Peters et al. 2009;
Cutignano et al. 2012; Carbone et al. 2014).
1 Psychrophiles as Sources for Bioinspiration … 9

Fig. 1.6 The morphology of Antarctic demosponges (a, b) is diverse. a Reprinted from Gutt
(2004c), photograph: J. Gutt ©AWI/MARUM (Germany), published under: Creative Commons
Attribution 3.0 Unported. b Reprinted from Gutt and Starmans (2004a), ©AWI/MARUM
(Germany), published under: Creative Commons Attribution 3.0 Unported

Interestingly, Antarctica has similar demosponge species richness as the Arctic—

352 versus 360 species, respectively (McClintock et al. 2005). Both demosponges and
hexactinellids habituating in Antarctic seas are important players of psychrophilic
benthic communities there (Little 1966; Dayton 1979, 1989; Barthel et al. 1991;
Barthel 1992a, b; Barthel and Gutt 1992; Barthel and Tendal 1994; Teixidó et al. 2006;
10 H. Ehrlich and A. Nikolaev

Fig. 1.7 Psychrophilic glass sponge Caulophacus arcticus (a) represents several levels (b, c,
d) of hierarchical organization

Gutt 2007; Dayton et al. 2013). We kindly recommend readers to find more infor-
mation in detail about diversity and biology of psychrophilic hexactinellids in the
chapter entitled “Cold Biosilicification in Metazoan: psychrophilic glass sponges”
prepared by Tabachnik and coauthors for this book.

1.7 Psychrophilic Biocalcifiers

Psychrophilic calcification occurs in both unicellular pro- and eukaryotic organisms

as well as in multicellular animals like bryozoans, corals, molluscs, crustaceans,
echinoderms, and fish. They are able to synthetize their calcium carbonate and/or
calcium phosphate-containing skeletons under frigid environmental conditions.
Similar to psychrophilic biosilicification, there is a lack of knowledge about
mechanisms of calcification at temperatures near the freezing point. Low temper-
atures lead to exponential decreases in chemical reaction rates (Margesin et al.
2008). Consequently, the thermodynamic aspects of biomineralization in the hard
tissues of psychrophilic biocalcifiers are of principal scientific interest.
Below, we take the liberty to give a brief overview on psychrophilic biocalcifiers
which belong to bacteria, yeast, fungi, foraminifera, bryozoans, corals, mollusks,
echinoderms, crustaceans and, finally, icefishes.
1 Psychrophiles as Sources for Bioinspiration … 11

1.8 Psychrophilic Microorganisms

The north and south polar sea ice environments are ideal habitats for studying
microbial psychrophilic communities from biochemical, ecological, and biogeo-
graphical points of view (for review see Morita 1975; Deming 2002; Thomas and
Dieckmann 2002; Ruisi et al. 2007). Some bacteria are psychrophiles that will be
additionally confirmed by their scientific names like Psychroserpens (Bowman
et al. 1997); Psychromonas antarcticus (Mountfort et al. 1998); Polaromonas,
Polaribacter, and Psychroflexus (Staley and Gosink 1999); Planococcus psy-
chrophilus (Reddy et al. 2002); Desulfotalea psychrophila (Rabus et al. 2004);
Psychrobacter cryohalolentis and Psychrobacter arcticus (Bakermans et al. 2006);
Psychromonas ingrahamii (Auman et al. 2006); Exiguobacterium sibiricum
(Rodrigues et al. 2006; Ponder et al. 2008); Flavobacterium glaciei (Zhang et al.
2006); Phaeobacter arcticus (Zhang et al. 2008); Sphingopyxis alaskensis (Ting
et al. 2010); Methanolobus psychrophilus (Chen et al. 2012); and Clostridium
psychrophilum (Perfumo et al. 2014). Not only bacteria, but archaea (Cavicchioli
2006), yeast (Di Menna 1960, 1966; Margaret 1966; Goto et al. 1969; de Garcia
et al. 2007; Xin and Zhou 2007; Margesin and Fell 2008; Lee et al. 2010; Turchetti
et al. 2011; Laich et al. 2014), and fungi (Richard et al. 1997; Robinson 2001;
Hoshino et al. 2003, 2010; Held et al. 2006; Singh et al. 2006; Kochkina et al. 2007;
Leung et al. 2011; Tsuji et al. 2013; Wang et al. 2015) can proliferate at 0–10 °C,
metabolize in snow and ice at −20 °C (Junge et al. 2004), and can survive at −45 °
C (Cavicchioli 2006). Beyond the problems of cryosurvival, interest in psy-
chrophilic microorganisms has increased recently because of their antifreeze protein
activity (Kawahara 2002; Cavicchioli et al. 2002; Gilbert et al. 2004) as well as due
to biotechnological potential of so-called “cold-adapted enzymes” for industrial
uses (Brenchley 1996; Russell 2000; Smalas et al. 2000; Feller and Gerday 2003;
Siddiqui and Cavicchioli 2006; Margesin and Feller 2010; Buzzini et al. 2012;
Feller 2013; Burhan et al. 2014; Alcaıno et al. 2015).
However, also some very intriguing findings on psychrophilic calcification in
bacteria have been reported. Thus, recently described gram-positive, aerobic bac-
terial isolates from a Canadian high Arctic permafrost active layer, known as
Planococcus halocryophilus strain Or1, habituate at temperatures as low as −15 °C
and high salinity of 18 % NaCl (Mykytczuk et al. 2012, 2013; Ronholm et al.
2015). It was observed that growth at −15 °C coincides with increasing
hydrophobicity of the cells and distinct extracellular encrustations closely associ-
ated with their cell walls. Researchers showed with strong evidence that the cell
envelope of bacteria grown at −15 °C is composed of 20 % calcium carbonate,
50 % peptidoglycan, and 29 % choline (Fig. 1.8) (Mykytczuk et al. 2016). It was
suggested that carbonic anhydrase, an enzyme responsible for biocalcification,
plays an important role in this unique case of psychrophilic biomineralization.
12 H. Ehrlich and A. Nikolaev

Fig. 1.8 Scanning and transmission electron micrographs of P. halocryophilus Or1 cells grown
at 25 °C (a–d), 5 °C (e–h), and −15 °C (i–p). Scale bars = 1 µm with exception of C, O, and P,
where scale bars = 500 nm. N nodular features, S sheet-like features, M peptidoglycan/murein,
PM plasma membrane. Reprinted from Mykytczuk et al. (2016) with permission of Springer

1.9 Psychrophilic Foraminifera

Mechanisms of biomineralization in foraminifera are still under investigations

(Hemleben et al. 1986; Erez 2003; de Nooijer et al. 2009, 2014; Robbins et al.
2016). These biocalcifying unicellular eukaryotes also habituate in cold water
environments worldwide (see for review Anderson 1975; Lagoe 1977; Alexander
and DeLaca 1987; Wollenburg and Mackensen 1988; Bernhard 1989; Bergsten
1994; Gooday et al. 1996; Hald and Korsun 1997; Suhr et al. 2008). For example,
foraminiferans such as Cibicides antarcticus and Adamussium colbecki are major
components of Antarctic ecosystems and play crucial roles in Antarctic carbonate
budgets. It was demonstrated that these species both contribute considerable
1 Psychrophiles as Sources for Bioinspiration … 13

amounts of CaCO3, potentially adding 5.94 × 109 kg ha−1 year−1 of CaCO3 to the
Ross Sea (Hancock et al. 2015).

1.10 Psychrophilic Sponges (Calcarea)

Calcareous sponges (class Calcarea) represent exclusively marine animals which

produce their skeletons of calcium carbonate. The morphology and microarchi-
tecture of calcitic spicules found in these sponges is very diverse (see for review
Haeckel 1872; Bidder 1898; Boury-Esnaultand Rützler 1997; Sethmann and
Wörheide 2008; Kopp et al. 2011).
Psychrophilic species of Calcarea have been reported from Arctic (Breitfuss
1896, 1898, 1933; Arnesen 1900; Tendal 1989; Rapp 2006, 2013) as well as from
Antarctic (Polejaeff 1883; Jenkin 1908; Dendy 1918; Brøndsted 1931; Rapp et al.
2011, 2013; Janussen and Rapp 2011; Rios and Cristobo 2014) seas. Some of them,
like Leucetta antarctica, are endemic. The specialized cells known as sclerocytes
are responsible for biosynthesis of calcitic spicular structures (Ilan et al. 1996) also
in psychrophilic calcareans; however, the mechanism of biomineralization in all
calcareous sponges is principally poorly understood. However, it is established that
α-carbonic anhydrases remain to be the key biomineralization enzymes in these
animals (see for detail Müller et al. 2013; Voigt et al. 2014).

1.11 Psychrophilic Bryozoa

Psychrophilic bryozoans are colonial benthic calcifying metazoans which have high
levels of Antarctic endemism (Schopf 1969; Hayward 1995; Peck et al. 1995;
Zabala et al. 1997; Rosso and Sanfilippo 2000; Barnes and Griffiths 2008).
Numerous reports have been dedicated to ecology, growth, feeding, and repro-
duction of these organisms living in cold waters (Winston 1983; Barnes and Clarke
1994; Barnes 1995, 2015; Figuerola et al. 2013). These studies are important also
from biomineralization view, as experimental results give answers to principal
questions about the rate of biocalcification at temperatures near freezing point of
water. For example, it was reported that the Antarctic bryozoan Melicerita obliqua
colonies form one segment per year, thus attaining a maximum length of about
200 mm within 50 years (Brey et al. 1998). Calcification in bryozoans appears to
be strongly genetically controlled. These invertebrates contain in their skeletal
structures calcite, aragonite, and mixtures of both (Smith 2014). Although the
principles of biomineralization in bryozoan have been recently reviewed in detail
(Taylor et al. 2015), there are gaps in our knowledge which need to be addressed
especially with respect to understanding of mechanisms of psychrophilic
calcification.
14 H. Ehrlich and A. Nikolaev

1.12 Psychrophilic Corals

Coral habitats span the range from tropical to polar areas including Antarctic seas.
Seventeen species of scleractinian corals occur in Antarctic waters (Fig. 1.9)
(Cairns 1982; Waller et al. 2008, 2011; Henry 2013). Despite the great interest and
numerous studies published on the calcification in corals, enormous gaps still exist
in our understanding of their psychrophilic biomineralization. The low temperature
tolerance of psychrophilic corals is well described (Fosså et al. 2002; Dodds et al.
2007; Roberts et al. 2009; Brooke et al. 2012). Some of psychrophilic scleractinians
are endemic species of the Antarctic region. For example, Flabellum impensum is a
very conspicuous species with large size and greater number of septa in comparison
to other species (Cairns 1990, 1995; Schejter and Bremec 2015). This coral is a
long-lived, slow-growing organism, with a low metabolic rate and a linear exten-
sion rate of approximately 1 mm per year, and a skeletal density that averaged
22 % more than the density of pure aragonite (Henry 2013).
Also, octocorals have been reported to habituate in Arctic (Cairns and Baco
2007) and Antarctic (Orejas et al. 2001, 2002, 2007; Taylor and Rogers 2015;
Dueñas et al. 2016) (Fig. 1.10) regions. One of the typical representatives of
psychrophilic octocorals is Primnoisis spicata that is related to family Isididae. The
axial skeleton of this species is very peculiar in that it contains both calcareous

Fig. 1.9 Heavily mineralized skeleton of Antarctic stony coral (Scleractinia, Cnidaria, Anthozoa)
1 Psychrophiles as Sources for Bioinspiration … 15

Fig. 1.10 Antarctic octocoral. Reprinted from Gutt (2004b), photograph: J. Gutt
©AWI/MARUM (Germany), published under: Creative Commons Attribution 3.0 Unported

(internode) and horny (gorgonin-based node) regions (Thomas and Mathew 1986).
Understanding of the principles of psychrophilic biomineralization especially in
these corals is of crucial interest because the biomimetic potential of Isididae
octocorals as models for bioinspired design of bone implants is well established
(Ehrlich et al. 2006a, b).

1.13 Psychrophilic Molluscs

In contrast to other biocalcifiers, molluscs have not been overlooked as biominer-

alogical resources, especially as model organisms for studies on shell and nacre
formation. However, we currently lack any information on the psychrophilic cal-
cification in corresponding mollusc’s species which represent numerous unique
phenomena. Some of them like the aragonite-shelled (thecosome) pteropod (sea
butterfly) Limacina helicina are even bipolar, occurring in both the Arctic and
16 H. Ehrlich and A. Nikolaev

Antarctic oceans (Comeau et al. 2009). The sub-Arctic bivalve Arctica islandica
can live in excess of 350 years (Schöne et al. 2005). The Antarctic cephalopod
mollusk Mesonychoteuthis hamiltoni is the largest known invertebrate (Rodhouse
and Clarke 1985; Allcock et al. 2001). The gizzard plates from the Antarctic
deepwater gastropod Scaphander cylindrellus contain the calcium oxalate dihydrate
mineral, weddellite (Lowenstam 1968), and not calcium carbonate.
The diversity of psychrophilic molluscs including endemic species like Antarctic
cephalopods Pareledone sp. (Allcock et al. 1997, 2005, 2007), Antarctic pteropod
Clione antarctica (Dymowska et al. 2012), Antarctic nudibranch Austrodoris ker-
guelensis (Diyabalanage et al. 2010), and the Antarctic clam Laternula elliptica
(Peck et al. 2002; Cummings et al. 2011) is well studied (Dell 1964; Linse 2003;
Ghiglione et al. 2013; Moreau et al. 2013). Recently, psychrophilic species L.
elliptica and L. helicina have been used as key indicators of the acidification
process in polar ecosystems (Morley et al. 2012; Bednaršek et al. 2014). L. elliptica
(Fig. 1.11) represents also a highly tractable mollusk model for studying psy-
chrophilic biomineralization (Sleight et al. 2015). Most attention, however, is paid
on transcriptome investigations using such species as Antarctic brooding gastropod
Margarella antarctica (Clark and Thorne 2015), Antarctic pteropod Limacina
helicina antarctica (Johnson and Hofmann 2016), and the Antarctic soft-shelled
clam, L. elliptica (Sleight et al. 2015; Park and Ahn do 2015).

Fig. 1.11 Antarctic clam L.

elliptica possess massive
proteinaceous matrix that
become visible after partial
decalcification. The role of
this organic matrix in
psychrophilic calcification is
still unknown
Another random document with
no related content on Scribd:
the Highlanders were scattered over the earth, to do what they
certainly have done well—a goodly share of the world’s work.
Now for Culloden! We—that is, four men of us—hired a horse,
driver, and carriage, and rode out to the desolate moor, which is
usually called “Culloden” by strangers and “Drummossie Moor” by
the natives. It is a tableland lying six miles northeast of Inverness
and not far from the Moray Firth. As we approached it, we could
discern the sunken lines of the trenches, in which about eighteen
hundred of the clansmen, killed in battle, were buried. In 1881, these
trenches of the different clans were marked by rough memorial
stones giving the clan names. At one part of the field was a stream
of water, to which the poor wounded wretches crawled to slake that
horrible thirst which comes so quickly to a soldier who has lost blood
and whose veins are drying up.
On one side was a cairn of stones about twenty feet high, reared
to mark the battle, in the front of which is set a tablet giving the
historical facts and date. But what touched us most deeply, as
Americans, was a colossal wreath of flowers and greenery hung
near the top. This token, though faded and its purple ribbons stained
by three months of summer rain and storm, told of “hands beyond
sea” and hearts that were saddened at the name of Culloden. I
asked who had hung that wreath upon the cairn and was told that it
had been sent by Scotsmen in America, whose ancestors had fallen
in that awful battle of April 16, 1746, in which the hopes of “Bonnie
Prince Charlie” were shattered and those of the House of Stuart to
reattain power came to an end. I understood that such a floral tribute
was offered annually.
 THE CAIRN AT CULLODEN
Some distance away was the place where the English cavalry
were held in reserve, to charge upon the fugitives and slaughter
them after they had broken and fled. Near the field also was a large
flat rock, which the Pretender had mounted to see the action and
scan its results. From this point of vantage, he fled, to suffer untold
hardships, while wandering for weeks, disguised as a woman, under
the care of the heroic Flora Macdonald. He was finally able to reach
the French ships, then lying off the coast for him, by which he was
able to get back to the Continent, there to end his days as a
drunkard.
Cumberland, the British general, knew that a failure to win on
this field, or a drawn battle, would mean a long-continued guerilla
warfare in the Highlands. So he gave orders to put to the sword all
the clansmen known to have been on the field. As we rode back to
Inverness, over which the English cavalry had thundered after the
battle, the intelligent driver pointed out more than one place, such as
blacksmith’s shops, rocks, and hollows, where fugitives had hidden
and whence they had been dragged out to be killed.
Culloden enables us to see what war was to the Highlanders,
what they meant by a campaign, and how far these men of the
claymore, broadsword, and target had advanced in military science.
The idea of these stalwart warriors, trained in clan feuds and
inheriting the prejudices and traditions handed down to them from
ancestors, was to go out in summer time, without special equipment,
commissary train, or dépôt of supplies. They would make a foray,
fight a battle or two, burn the enemy’s houses, drive off some cattle,
and then come home to divide the spoil—a system hardly higher in
dignity than that of the North American Indian highlanders, the
Iroquois.
The men of the glens cared little for firearms, whether musket or
cannon. Their favorite weapons from of old were the dirk and the
claymore. The latter was a long-handled, double-edged sword
weighing from five to seven pounds, with a handle often a foot long
and with one cross-bar for a hilt. This claymore, in which they
gloried, was a weapon quite different from the later single-edged and
basket-hilted sword, which did not come into use until well into the
eighteenth century. Their one idea of fighting was to make an onset
and come to close quarters. On their left arm they carried the target,
or round shield, made of light, tough wood, covered with bull’s hide,
stretched in one or more thicknesses and with boss or studs, and
sometimes furnished with a rim of metal, or armed with a sharp point
in the middle. With this defence, protecting more or less their faces
and body, they rushed upon the foe, in order to be free at once to
use, in older times, their claymores, or double-handed blades, or, in
later days, the broadsword in close combat. When fighting with
infantry armed with smooth-bore muskets and bayonets, they could,
after the first volley, fired at more or less close range, dash into the
files. Before the soldiers could reload, the Highlanders would be
upon them, dashing aside the bayonet thrust. Then, with stabbing or
cutting blow, the clansmen slaughtered their foes and thus made
firearms of little account.
 It is true that when large levies were made, as in the earlier
centuries, the Scottish spearmen were massed together and made a
formidable front, though as a rule, the English archers, with their
long-range missiles, were able to work havoc among the Scots, and
thus prevent them from getting into close hand-to-hand action. Thus,
the Southrons more than once ruined the chances and hopes of their
northern foes. In archery, the Scots never were able to compete with
the English.
Even when, later, some of the Highlanders possessed cannon,
they were apt to look with contempt upon anything which did not
permit them to charge in a rush and come to close quarters. In fact, it
was this unintelligent tenacity in holding on to a war equipment
which, even to the claymore, to say nothing of the target and
ordinary spear, had been discarded in other countries, that brought
the clans to final destruction at Culloden. On the Continent
improvements were made, first in favor of the pike and then of the
musket, with the dropping of anything like a shield, or defence, which
required the use of one hand and which could not resist a bullet. It
was a thorough knowledge of the Highlander’s conceit and
conservatism, which had become his weakness and was ultimately
to be his ruin, as well as the perception of the change in battle tactics
and the relative merits of bayonet and broadsword fighting, that
enabled the Duke of Cumberland, then only twenty-four years of
age, to win a decisive victory, such as older men of experience had
repeatedly tried to gain, but to no purpose.
Chambers wrote, in 1830, “The field of Culloden yet bears
witness to the carnage of which it was the scene. In the midst of its
black and blasted heath, various little eminences are to be seen
displaying a lively verdure, but too unequivocally expressive of the
dreadful chaos. They are so distinct and well defined that the eye
may almost, by their means, trace the position of the armies, or at
least discover where the fight was most warmly contested.”
The way toward Inverness, otherwise an unimproved, secondary
road, is fringed with many doleful memorials. There the daisy and
bluebell of Scotland have selected their abode, he tells us, as if
resolved to sentinel forever the last resting-place of their country’s
heroes. Not infrequently modern curiosity hunters have violated the
graves in order to secure some relic of the ill-fated warriors, to show
as a wonder in the halls of the Sassenach. The Gaels, with nobler
sentiments, have come more frequently to translate the bones of
their friends to consecrated ground afar, in their own dear glens of
the west. “But enough and more than enough yet remains to show
where Scotland fought her last battle and the latest examples of her
ancient chivalry fell to feed the eagles and to redeem the desert.”
Inverness in 1745, as Chambers describes it, was a royal burgh
in the vicinity of a half-civilized territory not yet emancipated from
feudal dominion. Though a seaport, it had only a slight local
commerce. The town bore every external mark of wretchedness. Its
people, even its shopkeepers, wore the Highland dress, in all its
squalor and scantiness; for the Highland plaids which we see to-day,
in silk and wool, and sold in shops of luxurious appointment, are
vastly different from the home-made fabrics of a century or more
ago. The Inverness people generally spoke Gaelic. A wheeled
vehicle had never yet been seen within the town, nor was there a
turnpike road within forty miles of its walls. Some contact by sea with
France and the dwelling in winter time of the Highland gentry in the
town shed some gleams of intelligence over the minds of the kilted
burghers. Yet when the Young Chevalier took up his residence at the
house of Lady Drummuir, hers was the only dwelling that had even
one room without a bed in it.
It was from Inverness that “Bonnie Prince Charlie,” in 1745,
marched out with his Highlanders to the gage of battle at Culloden,
of which we tell in another chapter. At neither of our two visits to the
Capital of the Highlands had we hosts or hostesses to invite us to
drink with them the inevitable cup of afternoon tea, without which a
Britisher does not feel that the island is safe, or that Britannia rules
the waves. So we must needs be satisfied with hotel service for our
Bohea and cups, though we are bound to say that the decoction was
excellent and the white-capped and snowy-aproned maid’s voice
was low and sweet.
As we chatted over our excursion to Drummossie Moor, we
recalled that the victor of Culloden, on arriving at Inverness, found
not only a considerable quantity of provisions, which had been
prepared for the poor Highlanders, but many of the Jacobite ladies,
who had attended their husbands during the campaign. They had
just enjoyed their afternoon tea-drinking and were preparing for an
evening ball, at which the Prince and his officers were to be
entertained, after his expected victory. It was the entrance of the
fugitives, who informed them of the fatal reverse their friends had
met with, which caused an abrupt change of plans.
Yet the lovers of the lost cause cease not their celebrations.
“Come o’er the stream, Charlie!” To this day, in the Highland glens,
one can hear old women singing to the tune of “Bonnie Prince
Charlie,” inviting him to “come over the border,” and feast himself on
“the red deer and the black steer,” promising, also, that his loyal
followers will “range on the heather, with bonnet and feather.” The
remnant of English Jacobites still drink to the health of the Stuarts
and hold an annual celebration in memoriam, in London and in
Philadelphia.
 CHAPTER XV
“BONNIE PRINCE CHARLIE”

Mary Queen of Scots, and Bonnie Prince Charlie! How they live
with us yet, casting their spell over the centuries!
If there is one figure in the past that still acts powerfully upon the
tradition, literature, and imagination of Scotland,—in a word, upon
that which remains and is imperishable, after stone and brass are
but mouldering relics,—it is the figure and fortunes of Charles, the
Young Pretender to the throne of Great Britain. With him ended
Celtic Scotland, Scottish feudalism, and the age of Highland
romance.
About the “Young Chevalier”—the image on the Scottish mind is
that of the fair youth in the full splendor of manhood; not the
wretched dregs of the human form that many years afterwards was
cast out of memory like an abominable branch. It is of the bonnie
young fellow that such songs as “Wae’s me for Prince Charlie,”
“Charlie is my darling,” “Come o’er the stream, Charlie,” and “The
White Cockade,” were written and are still sung. His full name was
Charles Edward Louis Philippe Casimir Stuart.
This young man, of extraordinary beauty and fascinating
manners, against the advice of his friends and most loyal supporters,
landed in Scotland, and summoning the Highland chiefs, who, by
affinities of blood, politics, and religion, were most attached to the
Stuart dynasty, asked for their support. One and all, they declared
against the uprising, but they, nevertheless, agreed to follow their
liege lord.
Born at Rome, on December 31, 1720, grandson of King James
II of England, and eldest son of James, the Old Pretender, who
called himself James III, Charles was nominated by his family the
Prince of Wales. Educated under brilliant tutors, he travelled through
Italy. He was able to speak English, French, and Italian, but could
never write well in English. Despite the previous failure, in 1715, of
his father, and the loss at sea by storm of a French fleet, with seven
thousand men who were to assist his Highlanders, Charles landed in
Scotland when most of the British army was in the Belgic
Netherlands. On August 19, 1745, in Glen Finnan, he unfurled his
standard as “James VIII of Scotland and III of England” against
George II and the Hanoverian dynasty of Great Britain. He wore the
Highland costume and won the hearts of the women by his charming
manners and manly beauty.
After a meteoric career, including a brilliant series of marches,
victories, occupation of Holyrood Palace in Edinburgh, invasion of
England almost to London, and sudden retreat, he had to face with
his loyal clansmen the King’s son William, Duke of Cumberland, with
an army specially trained to the use of the bayonet. The two forces
met on Drummossie Moor, near Culloden, April 16, 1746.
Cumberland’s men were in high spirits and fine condition, while the
ill-fed followers of Charles, hungry and weary after a night march,
numbered five thousand. His attempt to surprise the Duke and settle
the issue with cold steel had failed!
Against the advice of his officers, Charles ordered the battle.
After various manœuvres the armies faced each other for the bloody
decision, on which depended the fate of the House of Stuart, the
fortunes of the Highlanders, and the continuance of Scottish
feudalism.
One dreadful surprise awaited the clansmen. Cumberland,
trusting in the bayonet, had carefully drilled each of his men to have
the nerve to neglect the man striking at him with his broadsword, but
to stab at the fellow who, in expectation of dashing aside the bayonet
of the soldier in front of him, would expose his body to the oblique
thrust of his comrade on the right, duly fore-warned.
The day was one of chilly weather, with fitful winds and flurries of
snow. Early in the afternoon, the battle was opened by discharges of
cannon from the side of the rebels. But with this kind of work, the
men from the glens never were satisfied. Indeed, all firearms and
long-range weapons were unpopular with these brave fellows, who,
like Indians and semi-barbarians, enjoyed most that action which
was, as far as possible, independent and personal.
In several of their victories over the royal troops, as at
Prestonpans, for example, they had felt little or no annoyance from
the royal cannon, and had almost lost their fear of artillery.
Cumberland had nine thousand men and eighteen well-served
guns. Here, for the first time, the Highlanders were under heavy fire
of grape and round shot, to which they could not proportionately
reply. It is thought that if Charles at Culloden had let his swordsmen
rush at once upon the enemy the issue might have been different.
For half an hour the Duke’s cannon played effectively upon the
clansmen, who saw scores of their kinsmen stretched upon the
heath. After a few moments’ cannonade from their own side, and still
under the withering fire of the enemy’s heavy guns, the Highlanders
ranged themselves in masses, and according to their clans, made
ready for the terrific onset, which they supposed would decide the
battle. This it did, but not in the way they had hoped. It was the
Mackintoshes, who, unable any longer to brook the unavenged
slaughter of their comrades, broke from the centre of the line and
rushed forward through the smoke and snow to mingle with the
enemy. Yet the order to advance, though never delivered, had
already been given by Charles, the bearer being killed by a cannon
shot.
Cumberland’s troops, seeing the dark masses moving up the
slope, as in a great wave, stood in steady line. As the Highlanders
came to shock, the oblique thrust of the bayonets was a dreadful
surprise, for it prevented hundreds of clansmen from wielding their
favorite weapon, as most of them were thrust through before they
could swing their broadswords, or make the terrible double-handed
sweep with their claymores, on which they had counted. Soon the
moor of Drummossie had proved itself to be the valley of decision for
the hopes of the House of Stuart.
 Within two minutes the charge was general along the whole line.
Yet it was as if advancing into semi-darkness of whirling snow and
powder smoke. One survivor of the battle, a Highlander, said that
after rushing forward the first glimpse he received of the Duke’s
troops was, when the cloud of smoke and snow lifted, he saw the
white gaiters of the soldiers. The Duke’s cannon, now loaded with
grapeshot, and the musketry of his solid columns swept the field as
with a hailstorm. The three ranks in the front line of English Hessians
delivered simultaneous volleys, while the regiments of Wolfe—of
whom we Americans have heard in his later career at Quebec—
poured in a flank fire. Nevertheless, the right wing and centre of the
Highlanders fought with even more than usual gallantry and
resolution.
Notwithstanding the fact that they were outflanked, enfiladed,
and met by a heavy musketry fire in front of them, the right wing of
the Highlanders broke Barrel’s regimental front and passed the guns;
but their attack was checked by the bayonets of the second line.
Of the Highlanders who first rushed forward the majority were
hardly able to see their enemy for the smoke, until involved
inextricably among their weapons. Tn their onset, nearly all in the
front ranks fell before either bullets or the piercing weapons used
obliquely, as directed by the Duke, almost every bayonet being bent
or bloody with the strife. Nevertheless, the Highlanders, despite their
impending annihilation, kept on, line after line pushing forward, even
though only a few of those charging last reached the front files of the
royal troops. In parts of the plain, the dead lay three and four deep.
During all this time the Macdonalds, who, because their
ancestors at Bannockburn had fought on the right wing, had ever
afterwards, except on this occasion, occupied this position, would
not fight. They made no onset, and even received the fire of the
English regiments without flinching. They were dissatisfied because
they had been put on the left wing. At last, when the moment of
decision and defeat had come, there being no hope, they also fled
with the other clans.
 Charles had yet in reserve his foreign troops, and these, after
the mountaineers had been ruined, he hoped, as he looked on from
the mound at some distance off, would redeem the day. But though
there were instances of bravery among these men, yet, demoralized
by the wreck of the clans coming as fugitives among them, and
seeing the Duke’s army getting ready to charge with the cold steel,
they fled in a body. Thus the rout was complete. Charles, who had
made his last cast for a crown, seemed now unable to realize what
had happened. Confounded, bewildered, and in tears, he seemed
unable to act. His attendants were obliged to turn his horse’s head
and compel him to retreat, Sullivan his friend seizing the horse’s
bridle and dragging him away.
During the uprising of 1745–46, the local clans wore a red or
yellow cross or ribbon, in order to distinguish themselves from the
Stuart Highlanders, who were all dressed in about the same way,
except as to their bonnets. The Jacobites all wore the white cockade,
like that of the Bourbons of France, friends of the Stuarts. One of the
liveliest tunes played by the Highland pipers was “The White
Cockade.” It was the same air, with different words, which the fifers
and drummers of the Continental army played when the flag of the
Revolution was raised in the War of Independence. In fact, in looking
over the American musicians’ repertoire, from 1775 to 1783, one
might almost imagine that the chief music sounded under “the
Congress flag” of thirteen stripes and, after 1777, under “Old Glory”
of later Revolutionary days, was Scottish. Even the strains of
mournful music, over the graves of the slain American patriots, was
“Roslyn Castle.”
One fifth of the Highland army was lost at Culloden. Of the five
regiments which charged the English, almost all the leaders and
front rank men were slain. These numbered nearly a thousand in all.
The actual battle lasted about forty minutes, much of it in distant
firing; but the charge and the crossing of the cold steel were all over
in a quarter of an hour. The number of killed, wounded, and missing
of the royal army was three hundred and ten. The victory was mainly
attributable to the effect of the artillery and musketry of the royalists;
but in Munro’s and Barrel’s regiments, many of the soldiers put to
death one, two, or more Highlanders each, with their bayonets, and
several of the dragoons, sent in pursuit, were known to have cut
down ten or twelve fugitives each in the pursuit.
 CHAPTER XVI
THE OLD HIGHLANDS AND THEIR
INHABITANTS

The Highlands, geologically speaking, is an island of crystalline

rock set in a great sea of younger formations. The great glen which
forms the trough of the Caledonian Canal is a mighty earth rift. When
once across this line of rock and water, we were in the Highlands. In
one summer visit, we spent a part of our vacation at Crieff, which lies
at the base of the Grampian Hills and at the entrance to the
Highlands. Here the beauty, fashion, and intelligence of the United
Kingdom in August gather together. What was once a “hydro,” but is
now a fine hotel, was crowded to its utmost capacity. In the
evenings, entertainments of music, with dancing and recitations by
the young people, were enjoyed. In the mornings, we took horses
and carriages and drove through many leagues of the lovely
scenery. At another time, in a later year, the automobile served us
while glancing at a hundred linear and many more square miles of
Scotland’s glory.
Yet every time we were in the Highlands and in whatever shire,
the old song, learned in childhood, came to mind—“O where, tell me
where, has my Highland laddie gone?” Ross and Cromarty, now
united in one and the largest of all the counties in Scotland, is the
most thinly populated of all. In fact this great area has been
“improved” by its landed proprietors promoting the emigration of its
former inhabitants. There is only a fraction left of the Highlanders.
The Celtic element is but a survival, a remnant, and the Gaelic
tongue is like a flickering flame, almost ready to die out.
What is the reason? Is it, in part at least, because nature is so
niggardly? Again, is it not true that “those who take up the sword
shall perish by the sword”? Did the traditional Highlands and
Highlanders exist, or gain their place in romance and history, chiefly
through the human imagination?
Scottish history and poetry show that originally, even as a
swordsman and fighter, the Highlander possessed no special
superiority over the Lowlander, but in the seventeenth century, as in
the modern days, which we of ’61, as well as of 1915, remember,
and have seen demonstrated, the best prepared people, to whom
arms are habitual, and to whom military training is a personal
accomplishment, will, at the first beginning of war, at least, be pretty
sure to get the advantage. In a prolonged struggle, it is resources
that tell. Wars are not ended by battle, but by manifest reserves, with
power to follow up victory.
It was western Scotland, of azoic rock, a far-off corner of Europe,
that had the singular fortune of sheltering the last vestiges of the
Celts—that early race of people who, once placed upon the centre of
the ancient continent, were gradually driven to its western
extremities.
A notion, held tenaciously by the Highlanders, was that the
Lowlands had originally been their birthright. Many of them practised
a regular system of reprisal upon the frontier of that civilized region,
with as good a conscience as a Levant pirate crossed himself and
vowed to burn candles of gratitude before the Virgin’s picture, if
successful in robbery. To maintain this philosophy and practice, the
use of arms was habitual and necessary among the Highlanders.
While among the Lowlanders cattle-lifting and other methods of
rapine were considered as the business of thieves and scoundrels, it
was usually reckoned by the Highlanders to be an eminently
honorable occupation, partaking of the prestige of a profession. How
finely does Sir Walter Scott bring out this sentiment, when Roderick
Dhu answers Fitz-James, who charges the Highland chieftain with
leading a robber life.
Moreover, what still tended to induce military habits among the
Gaelic mountain folk, and what still maintains most wars, in the same
spirit, though on a larger scale,—national instead of private,—was
the hereditary enmity against each other, systematically maintained,
purposely cultivated and instilled in their children. In what respect
were the clan feuds and fights of the Celtic Scots any nobler than
those which so long distracted China, Japan, and Iroquois and
Algonquin America? With such philosophy dominant as still in our
day creates armies and navies, while being no more ethically worthy,
it was required that every man capable of bearing arms should be in
perpetual readiness to foment war, or to seize or repel opportunities
of vengeance. In fact, the hideous brutality of Confucian, Japanese,
Iroquois, Scottish, and Albanian codes of vengeance alike befitted
the common savagery that runs counter to the teachings of the
Universal Man of Nazareth.
The Celtic Highlanders were nominally subjugated by the iron
hand of Cromwell. Of this mighty man, Dr. Johnson says, “No faction
in Scotland loved the name of Cromwell or continued his fame.
Cromwell introduced, by useful violence, the arts of peace. People
learned to make shoes and plant kail.” Shoes were not common in
this part of Scotland until as late as 1773.
At the Restoration of the Stuarts, in the person of Charles II, the
Highlanders, with no illustrious and stimulating example before them,
rebounded into all their former privileges and vigor. They were kept
in arms during the reign of the last two monarchs, who fomented
those unhappy struggles, on account of religion, which have made
the Stuart name so detested. The patriarchal system of laws, upon
which Highland society was constituted, disposed these
mountaineers to look upon these unhappy princes, Charles I and
James II, and upon the Pretenders, who came after them, as the
general fathers or chiefs of the nation, whose natural and
unquestionable power had been wickedly disputed by their rebellious
children. Hence at Killiecrankie, Prestonpans, Falkirk, and Culloden,
they fought with the same ardor that would induce a man of
humanity to ward off the blow which an unnatural son had aimed at a
parent. In a word, as to political education, they had only the ideas of
feudalism in which they were steeped.
Having myself lived under feudal institutions, and seen the daily
workings of a society, graded from lowest to highest, although with
many variations, and fixed in customs which seemed to me to be
tedious, absurd, and ridiculous, as well as interesting and
fascinating, and living meanwhile under the shadow of castle walls
and towers, crossing daily the drawbridge and often visiting the
towers of the citadel, I could understand the mediæval processes of
thought, so long surviving in western Scotland. I was able to
appreciate also these Scottish castles, whether still maintained as of
old, intact and modernized, or in ruins, and easily re-create in
imagination the mental atmosphere and customs of the old feudal
days, when swords were an article of daily dress and frequent use,
and the steel blade the chief bond and instrument of social order.
The border ruffianism of “bleeding Kansas” in the West and much of
the old social situation down South, in cotton land,—the pride and
contempt on the one side and the hatred, with occasional cattle-
lifting propensities, on the other, especially in the Southern
Highlands,—of which in my boyhood I heard so much, helped me to
enjoy not only Scottish history, but Sir Walter Scott’s inimitable word
pictures in prose and verse. One can describe most of the
spectacular phenomena of Japanese as well as Scottish feudalism in
Scott’s verse and prose. His writings make illuminating commentary.
It was hard for the Lowlanders, after their discipline under the
feudal system had passed with the institution, to understand or get
along peaceably with the Highlanders, who hated industrialism,
shop-keeping, and money-making. Highland poverty and rawness
are in the main the immediate inheritances, even as the old semi-
civilized life was the direct result, of feudalism. The reason why the
dwellings of the plain people in the rocky regions were, even in our
day, so wretchedly poor and bare, is revealed in the book of Mair,
entitled “De Gestis,” published in Latin in 1518, concerning land
tenure. He says: “In Scotland the houses of the peasants are mere
small thatched huts, and the cause is, that they do not hold their land
in perpetuity, but only rent on a lease of four or five years at the will
of the lord; therefore, though there are plenty of stones, they will not
build neat houses, nor will they plant trees, or hedges to the woods,
nor will they enrich the soil; and this is to the no small loss and
disgrace of the whole realm. If the lords would give them their land in
perpetuity, they would get double or triple the money they now have,
because the peasants would cultivate the land incomparably better.”
This system of land tenure, which in theory and practice made
the laird the landowner and the tenant, or worker of the soil, a virtual
serf or semi-slave, sufficiently indicates the grounds and nature of
the Highland chief’s power and the degradation of the average or
common man. In almost every clan, there were subordinate chiefs,
cadets of the principal family, that had acquired a territory and
founded separate septs. In this community, the majority of
commoners were distinct from the “gentlemen,” who were persons
who could clearly trace their derivation from the chiefs of former
times and assert their kinsmanship to the present one. Below this
clan aristocracy were the mass of plain fellows (“kerns”) who could
not tell how or why they came to belong to the clan and who were
always distinctly inferiors.
There were several distinctions, based on ability, of status and
condition. The commoners were little better than serfs, having no
certain idea of a noble ancestry to nerve their exertions or to purify
their conduct. It was not to these, but to the gentry, that the chief
looked for active service and upon whom he depended in time of
war. These upper grades of men did most of the fighting, while the
larger body of common retainers (“kerns”) were left behind, during a
raid, to perform the humbler duties of driving the cows or tilling the
fields. Or, if they accompanied the foray, they were put in the rear
ranks and given poor arms, sometimes being provided only with
dirks. To illustrate these facts there were and are many stories told
and traditions handed down. Note the incident in “The Lady of the
Lake”:

“Because a wretched kern ye slew,

Homage to name to Roderick Dhu?”

In a word, in Scotland and in Japan, of which we can bear

witness from personal experience, social evolution among clansmen
and arms-bearing men had begun and continued, though separation
had early taken place between the fighters and the field laborers. In
both countries the process and result were much the same.
Moreover, after the Reformation, the proud Highlanders, clinging to
the old faith and traditions, looked down, with even greater contempt
than before, upon the commercial Presbyterians of the Low
Countries. They regarded with absolute horror the newer social and
political order, which in their eyes was but a dark system of
Parliamentary corruption. They were only too ready to believe the
stories of luxury, extravagance, and predatory dishonesty, which
were supposed to be rife and chronic in London. Here, too, human
nature, Japanese and Scotch, was as much alike as in a pair of
twins, born of the same mother, and throughout history running in
parallel lines of action.
Moreover, in both Scotland and Japan, it was the bayonet
against the sword. The men of mediæval mind in both countries
wore and wielded blades and looked upon the use of firearms as
something mean and cowardly. Believing, to the last, in the rush
against uniformed men in ranks and in slashing with two-handed
sword strokes (the Japanese swordsmen using a mat shield, where
the Highlander employed a target), both Scot and Nipponese met
failure against the triangular stabbing tools that ended feudalism. In
Tokio, the bayonet monument on Kudan Hill tells a story. Here,
history is told in steel.
What did more than anything else to open the Highlands and
break up the very idea of a “hermit nation” was a system of roads
which was carried out mainly during the sixteen years between 1726
and 1742, by the British field marshal, George Wade. Though born in
Ireland (whence also came the great soldier and diplomatist, Wade,
of China), he knew well the Gaels of both the island and the
mainland. He spent two years studying the problems of the
Highlands, economic and social. He had had long service with the
army in the Belgic Netherlands, Portugal, Spain, and the
Mediterranean Islands. During the Jacobite outbreak of 1715, he
acted effectively as military governor. Having later again made a
thorough study of the Highlands and their inhabitants, he was made
commander-in-chief, in order to give effect to his own
recommendations. He cut roads through the most important strategic
places and lines of country. In the course of this engineering work he
superintended the construction of no fewer than forty stone bridges.
It is this road-making which constitutes his chief title to fame, as the
old distich intimates:—

“Had you seen these roads before they were made,

You would lift up your hands and bless General Wade.”

In a word, he made possible the pacification of the Highlands, by

a system of hard-faced or “metalled” roads. Dr. Johnson, who saw
the results of Wade’s peaceful campaign, when the work was fresh
and the results novel, is unstinted in praise of Wade. In fact, it is
quite probable that, except for these new highways, the great man’s
“Journey to the Western Islands of Scotland,” in 1773, would not,
perhaps could not, have been taken.
The houses the Highlanders of a century ago lived in are
described by Dr. Johnson. The construction of a hut, he tells us, is of
loose stones, arranged for the most part with some tendency to
circularity and placed where the wind cannot act upon it with
violence, and where the water would run easily away, because it has
no floor but the naked ground. The wall, which is commonly about
six feet high, declines from the perpendicular a little inward. Some
rafters are raised for a roof, which makes a strong and warm thatch,
kept from flying off by ropes of twisted heather, of which the ends,
reaching from the centre of the thatch to the top of the wall, are held
firm by the weight of a large stone. No light is admitted, but at the
entrance and through a hole in the thatch, which gives vent to the
smoke. The hole is not directly over the fire, lest the rain should
extinguish it, and the smoke therefore fills the place before it
escapes.
 THE SCOTCH BRIGADE MEMORIAL
Entering one of this better class of huts, Dr. Johnson found an
old woman whose husband was eighty years old. She knew little
English, but he had interpreters at hand. She had five children still at
home and others who had gone away. One youth had gone to
Inverness to buy meal—by which oatmeal is always meant. She was
mistress of sixty goats and many kids were in the enclosure. She
had also some poultry, a potato garden, and four shucks containing
each twelve sheaves of barley. Huts in building and equipment are
not more uniform than are palaces, and hers was divided into
several apartments. She was boiling goat’s flesh in the kettle for the
next meal. With true pastoral hospitality, she invited her guest to sit
down and drink whiskey. Sweetening was obtained from honey.
Probably the reason why marmalade is so much used by the modern
Scots is because of old their ancestors used a great deal of honey,
of which marmalade, usually made from oranges imported from
Spain, takes the place.