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CONTENTS

Title Page
Introduction
Life
Inheritance
The cell
DNA
Sex
Evolution
Classification
Ecology
Animal relationships
Neo-Darwinism
Origins of Life
Humans and genetics
Genetics and technology
Glossary
Picture credits
Copyright
About the Author
Other Titles in the Series
About the Book
Introduction

I n its simplest terms, genetics is the study of inheritance. However,

looking a little deeper, there is nothing simple about it. Genetics tells
us how a body can grow from a single cell; it shows how life on Earth
has changed in a myriad ways over billions of years; and it forms a
central plank in the fight against disease. What’s more, it also has the
potential to create new technology that will transform society,
ensuring health for all and perhaps even allowing us to control the
future development of our species and reshape the living world.

As a science, genetics is relatively new: its foundations date from the

1850s, but those many different strands were not drawn into a single
field until the early 20th century. It was slow going at first, and not
until the 1950s did the great mysteries of genetics begin to give up
their meanings. First was the discovery of the DNA double helix, and
after that the so-called ‘Central Dogma’, which shows how an
inanimate chemical code can result in a living body. Progress
accelerated rapidly as we unlocked more of the secrets of the gene,
but even today, despite huge advances, there are many riddles within
our DNA that we are still to solve. We may have learned how to
decipher the genetic code, but the work of translating what it all
means is still proceeding.

Genetics draws from many fields, such as chemistry, biology,

agriculture, engineering, even information theory and statistics. For
many, the expectation is that genetics can tell us exactly who we are,
what’s ‘in the genes’. Long before the science of genetics existed, our
ancestors would have understood that a child was a unique blend of
characteristics inherited from its parents. However, the extent to
which the nature of our genetic code rules our behaviours and
personalities is proving the most difficult puzzle to solve. Perhaps the
latest interests of genetics, such as stem cell research, epigenetics
and artificial biology, will provide those missing pieces – certainly
these intriguing areas of research suggest that genetics will continue
to have a huge influence on medicine and our understanding of what
it means to be human in the 21st century and beyond.
Life

W hat is life? In a nutshell, scientists would define it as a self-

replicating process that requires at least one ‘thermodynamic
cycle’. To put that another way, something that is alive is able to
make a copy of itself, and it does this by harnessing a source of
energy, using it to transform chemical resources in some way. The
supply of energy must be continuous; if the energy source were to
become unavailable, or the life form became unable to tap it, then
the result would be death. That is something else unique that life can
do: it can die.

According to this definition, the simplest life form is a strand of

nucleic acid, something like RNA (see here). This chemical is able to
use its own molecule as a template for a copy of itself. However,
such a life is incredibly precarious, and over billions of years of
evolution, a multitude of life forms have developed abilities that
ensure survival. These abilities are set out in genes, and they govern
the success or failure of a life. To understand life, one must begin
with genetics.
Types of organism

T he number of different types, or species, of organism on Earth is

estimated to be anywhere between 3 and 30 million, with most
biologists erring towards about 9 million.

The simplest and oldest life forms are the bacteria, which have a
body made from a single tiny ‘prokaryotic’ cell (see here). They are
joined by the archaea, which to the uninitiated look more or less the
same but have some important distinctions. Other single-celled
organisms, including things like amoebae and protozoa, have much
larger and more complex cells, and this ‘eukaryotic’ cell type (see
here) is the one used by multicellular organisms such as plants,
animals and fungi.

Every species of organism has a unique way of life, but members of

any biological group share more characteristics with each other than
with the members of other groups. However, all life forms share a set
of abilities: they sense the surroundings, excrete, reproduce, grow,
respire and require nutrition.
Metabolism

T he broad term ‘metabolism’ encapsulates the link between

chemical activity and life – the myriad chemical processes that are
occurring inside every organism are described as its metabolism. In
very general terms, these include the way the organism handles its
energy supply, and how it uses this to grow and repair its body,
making use of simple chemical building blocks.

Metabolic processes fall into two general types: anabolism and

catabolism. The former involve building larger, more complex and
more ordered structures out of smaller units. (That is why a sports
cheat might use an ‘anabolic steroid’, a chemical that builds muscle.)
Catabolism, in contrast, involves splitting large structures into smaller
ones (this includes processing unwanted waste materials to generate
energy). Anabolic and catabolic processes are constantly working
together to release manageable packets of energy and then put them
to work in keeping the organism alive.
Feeding

E very living thing must feed, or putting it more precisely, they must
access a source of nutrition. Plants get this in the form of sugars
from photosynthesis and mineral nutrients absorbed from their
surroundings (soil is a good place to start). Animals and fungi get their
nutrition from the bodies of other organisms. Some single-celled
organisms can get nutrition using both techniques!

Nutrition has two main purposes. First, it is a source of chemical

energy that can be extracted and put to work in the body (the best
examples of this are glucose and other sugars). The second purpose
is as a stockpile of the raw ingredients required to build a body. The
requirements of different organisms vary wildly: plants are able to
build everything they need from water, carbon dioxide, and a menu of
minerals such as nitrates and phosphates, while animals need more
complex nutrition, such as fats, starches, proteins and a range of
crucial helper chemicals, known collectively as ‘vitamins’.
Respiration

W hen most people hear the term ‘respiration’, they tend to

assume it relates to breathing. But while this is indeed the
word’s common medical context, biology gives it a wider meaning: in
fact, all organisms respire, whether or not they breathe in and out in
the way that vertebrate animals do.

Biologically, respiration is defined as the metabolic process that

releases energy from sugar or other chemical fuels. Typically, this
involves the fuel molecules being oxidized – exactly the same
chemical reaction involved when materials combust in air. The
respiration of glucose, one of the most common sugars, for example,
can be written in the form of a chemical equation as shown opposite.
This demonstrates that glucose reacts with oxygen to produce carbon
dioxide and water, plus some energy. If raw glucose is burnt in air,
the reaction produces flames and heat, but within a living cell it can
be heavily regulated, allowing small packets of energy to be released
in several steps.
Photosynthesis

A s the word suggests, ‘photosynthesis’ is the process of ‘making

with light’, and the end product in question is glucose sugar.
Photosynthesis takes place in the leaves and the other green parts of
plants and other photosynthetic organisms. The colour is important
because the energy from sunlight is absorbed by a pigment chemical
called chlorophyll in the plant’s cells – chlorophyll itself appears green
because it traps the blue and red wavelengths of sunlight while
reflecting other colours.

Chemically, photosynthesis is the reverse of respiration, with carbon

dioxide and water molecules being combined to make glucose
molecules and oxygen, all powered by the energy channelled from the
chlorophyll molecules. While carbon dioxide is the waste product of
respiration, photosynthetic organisms produce waste oxygen, which
is released into the air. Nearly all of the oxygen in Earth’s atmosphere
(about 20 per cent of all the air) originated as the by-product of
photosynthesis.
Growth

T o qualify as living, an organism needs to be able to grow – at least

at some point in its life. For most complex organisms this is a
simple thing to verify. The majority of multicellular life forms – those
with bodies of more than one cell – grow from a single cell into an
embryo and on to a fully developed adult. This growth is achieved by
the division of cells (see here and here), with every cell in the body
being descended by some route from that first single cell, known as
the zygote.

The growth of single-celled organisms – things like bacteria and

amoebae – is less clear cut. They too can divide their cells, but
instead of creating a larger body, they produce a new and
independent individual. In these cases, growth and reproduction are
two sides of the same coin. Therefore, the best definition of growth is
the ability to produce new cells from older cells. This is the concept
that lies at the heart of cell theory (see here), a central tenet of life
science.
Reproduction

I t could be said that the primary goal of an organism is to survive.

However, that survival is really a means to an end – all organisms
are striving to make a copy of themselves or something close to it. In
other words the true purpose of biological life is reproduction. There
are many modes of reproduction, ranging from organisms simply
dividing in two to a complex process of courtship, mate selection and
parental care. However, broadly speaking there are two types of
reproduction: sexual and asexual. The former involves two parents
and the latter requires only one (see here and here).

The struggle to survive and reproduce is the driving force behind

evolution by natural selection (see here), the process that shapes the
millions of species that live on Earth. However, this evolution is a by-
product of reproduction. The genetic purpose of reproduction is to
make new copies, and many of them, of the DNA molecules in all
bodies, reproducing the information that we call genes.
Excretion

J ust as an organism takes in nutrients and other raw materials

from its surroundings, it must also remove the waste products of
metabolism – a process known as excretion. Despite common usage,
the voiding of the bowel, passing faecal matter, out of the body is not
actually excretion in biological terms: instead, it is defecation or
egestion. The crucial difference is that the unused food has not really
entered the body – it has only passed through the gut, a hollow tube
that runs through the body. True excretion is the process of taking
waste products – which may be harmful if left to accrue – from the
body’s tissues and expelling them.

In human biology the chief mode of excretion is urination, whereby

excess water and nitrogen-rich waste in the form of urea are
released. Excretion can also occur directly through the skin as
sweating. In addition, the release of carbon dioxide generated by
respiration processes is also a form of excretion.
Senses

A ll life forms are able to detect changes in their surroundings and

respond to them. For single-celled organisms this may be simply
a matter of detecting a chemical change, such as the salinity of water
or the presence of nutrients or toxins. Plants, meanwhile, are
sensitive to light, gravity and sometimes pressure – they grow
towards light and away from the pull of gravity, and some adjust their
growth patterns to wrap themselves around other objects they
contact.

Animal senses are much more advanced, befitting their active

lifestyles. The five used by humans are somewhat ubiquitous:
hearing, smell, taste, vision and touch. The last of these is a complex
mix of detectors on the body surface, sensitive to heat, cold,
vibrations and pressure. Other animals can sense things beyond a
human’s abilities. Many insects and other arthropods can detect
ultraviolet light; sharks and their cousins can detect electrical activity
in another body, while many other animals appear to sense Earth’s
magnetic field.
Inheritance

T he science of genetics is relatively new. Its first steps were made

in the 1850s and the term ‘genetics’ was not coined until 1905. It
was, however, a new word for an old field of enquiry: inheritance.
Since prehistoric times it was well understood that children inherited
some of the attributes of their parents. Characteristics such as hair
colour, face shape and height are passed on in families, from
generation to generation. This applies as much to animals and plants
– especially those used in farming – as it does to humans.

The search for the mechanisms of inheritance led to the science of

genetics and the theory of evolution, but it did not begin there. The
ancient Greek theory was ‘pangenesis’, which proposed that every
body part sent information via the semen and menstrual blood to
create a tiny person, or homunculus, that grew inside the mother.
Charles Darwin himself espoused something like this, saying inherited
traits travelled between generations as a swarm of tiny packets called
‘gemmules’.
The gene

T he term ‘gene’ was coined in 1909 by the Danish botanist Wilhelm

Johannsen. Its roots lie in the word ‘genesis’ meaning origin.
Charles Darwin and his colleagues in the late 1800s referred to a still-
hypothetical ‘genetic’ material that transmitted inherited traits. The
study of that process became known as genetics in 1905 (thanks to
English biologist William Bateson), and soon after Johannsen
introduced the concept of the gene.

Johannsen had no idea what form genes took. His term simply meant
a unit of inheritance: the genes inherited from the parent carry the
instructions required to build the body of a child. The term is also
used to describe particular measurable characteristics, so there is a
gene for hair type, eye colour, etc. However, today we know that
genetic material is a code-carrying molecule of DNA, so a section of
DNA can also be described as a gene. Matching this chemical
definition of genes with the anatomical one is a key goal of genetic
research.
T he core activity of genetics is to identify genes among the DNA held in cells, and
figure out their function.
Gregor Mendel

P erhaps surprisingly, the founding figure of genetics was a

German-speaking monk, living in the northern reaches of the
Austro-Hungarian Empire in the mid-19th century. Gregor Mendel’s
work, carried out in the cloistered garden of the Abbey of St Thomas
in Brno (now a Czech city), was completely ignored from its
publication in 1866 to the start of the 20th century, but nevertheless it
contained the basic tenets of genetics that still apply today.

Mendel (1822–84) made his discoveries through experiments breeding

pea plants in his garden. He had no knowledge of DNA, referred little
to cell biology and, instead of the term ‘gene’, used the word ‘factor’.
However, Mendel was able to glean some universal rules of genetics
from the way the different characteristics of the pea plants were
passed from generation to generation. These fundamental rules are
the foundations of the core inheritance process, which is called
Mendelian genetics in his honour.
Mendel’s crosses

G regor Mendel made his discoveries by diligently controlling which

pea plants were allowed to breed with which others. He was
aided in this endeavour by the fact that peas can self-cross, meaning
a plant can use its own pollen to produce seeds.

Mendel identified several inherited traits, such as flower colour or

shape and plant height. He worked on all these traits, but taking
height as our exemplar, Mendel isolated a tall plant that always
produced tall daughter plants when crossed with itself, and a short
plant that always produced short offspring. He then cross-pollinated
these two plants to produce offspring (seeds) with one tall and one
short parent. He found the first generation of offspring grew into tall
plants. Next he self-crossed one plant from the new generation.
Three quarters of its offspring were tall, a quarter were short. The
same thing happened for all the traits he tested. Mendel’s theories of
inheritance were deduced from these startlingly consistent results.
Mendel’s Laws

U sing the results from his many thousands of breeding

experiments carried out over several years, Gregor Mendel
outlined what he saw as universal truths about inheritance.

Mendel’s ‘Law of Segregation’ said that each plant had two versions
of each factor (gene). When it came to making pollen, the paired
versions of each factor were always split. Any offspring would inherit
only one version from each parent, with the two combining making a
new pair. Another rule, the ‘Law of Independent Assortment’, states
that every factor moves between generations independently of the
others. A third law, the ‘Law of Dominance’ asserts that some types
of factor have a hierarchy that leads to dominant ones being
expressed in the organism’s outward appearance, while recessive
ones remain hidden. Later research would come to qualify the second
law, and some regard the third as less significant because it does not
apply to all factors, but together these laws have become the
foundation stones of classical genetics.
Mendel’s diligent experiments with pea plants gave the first insight into how inherited
factors controlled development.
Phenotype

C lassical genetics draws a line between our two definitions of a

gene: a gene can be understood as a chemical entity – a piece of
DNA – or as an inherited trait, anatomical or otherwise. Mendel’s
discoveries showed that the two concepts were not interchangeable.
To illustrate this, geneticists invented the term ‘phenotype’.

The phenotype is the outwardly expressed end result of the genes

that are inherited. It is the tallness of the pea plant, the colour of your
hair or the body plan of an insect. It can also relate to animal
behaviours (sometimes referred to as the ‘extended phenotype’).
There is often a degree of learning involved in behaviours, such as
migration, hunting and nest building, but they are nevertheless
ultimately inherited from the parents. Mendel’s master stroke was to
figure out the link between the phenotype and the way genetic
material is transferred. That genetic material has been given another
name: the ‘genotype’.
Genotype

A n organism’s genotype could also be simply described as its

genetic make-up. It is a description of the various genes inherited
from its parents. As Gregor Mendel discovered, all organisms get one
version of each gene from each of their parents, and so the genotype
is made up of these pairs.

A particular genotype does not automatically lead to a related

phenotype. In fact, the same phenotype – for example, the tallness of
a pea plant – can result from a set of different genotypes (albeit a
small set). The mechanisms at play are twofold. Firstly, the different
versions of the gene interact and combine with each other in
particular ways – described by the ideas of genetic dominance (see
here) and Mendel’s Third Law. Secondly, the environment in which
the organism finds itself also has an impact on how it grows and
develops, by varying degrees from gene to gene.
Alleles

T his unusual sounding word derives from German and means

something like, ‘of one another’. It is however a very useful term
in genetics: an allele is one of several possible versions of a gene. So
using the example of Mendel’s pea plants, the gene for plant height
has two alleles: tall or short. Another example is eye colour: blue,
green, grey, brown and hazel are best described as alleles of the
same gene.

A genotype contains two alleles for each gene. If the alleles are
identical, then it is described as homozygous – in other words, when it
comes to dividing the alleles up to make the sex cells that are used in
producing the next generation (pollen, sperm, eggs etc), each cell will
definitely contain the same allele. When a genotype contains two
differing alleles it is described as heterozygous. As a result, half the
sex cells will have one allele, and half the other. Nevertheless,
homozygous and heterozygous genotypes can still produce the same
phenotype, thanks to an additional complication known as dominance
(see here).
T hese guinea pigs all have the same gene that controls hair colour; but each has
inherited a different version, or allele, of that gene.
Genome and gene pool

A genome is the total genetic material used by an organism. The

concept of our own human genome has become familiar ever
since the launch of the Human Genome Project in 1990. That effort to
map all of the genetic material used to make a human body was
completed in 2003, although the work continues, figuring out how
that material is divided up into somewhere between 20,000 and
25,000 genes.

Other organisms with mapped genomes include the E. coli baterium,

the worm Caenorhabditis elegans and the fruit fly. The amount of
genetic material – the DNA – in each organism varies, as do the
number of genes.

The ‘gene pool’ is another familiar term but one with a rather
different meaning. It refers to the enire set of genes, including their
many different alleles, that exists throughout a population of
organisms. The gene pool represents the genetic variation wihthin a
group of organisms.
As well as studying the genes of an individual, geneticists also seek to understand the
behaviours of genes shared by a community, population or entire species.
Hybrids

In general terminology, a ‘hybrid’ is an organism that is the product

of a cross between two distinct breeds. In terms of biology, and
genetics in particular, however, a hybrid is an organism that has a
heterogametic genotype. Put more simply, the organism has inherited
two differing alleles, or versions of a gene, from its parents.

Gregor Mendel’s success with solving the puzzle of inheritance

involved repeated hybridizations. Although he had no idea of how it
was happening, Mendel correctly surmised that his crosses of pea
plants were creating hybrids – pea plants that had inherited two
different versions of a gene.

It was this breakthrough that allowed him to discover that one allele is
not always equal to another. Some are dominant over others, and it is
this interplay of alleles that shows how a particular genotype leads to
a corresponding phenotype.
T his żubroń is a hybrid of domestic cattle and European bison.
Dominance

G enetic dominance is the overriding feature that explains the

results of Mendel’s hybrid experiments (see here). Returning to
the example of a tall pea plant crossed with a short one, the tall
parent has the genotype TT, with T being the tall allele. The short
parent’s genotype is tt, with t being the short allele.

The next generation of plants all receive a T from one parent and a t
from the other. They all have a genotype of Tt. The T allele is
dominant over the t, and so all Tt genotypes lead to a tall phenotype.
Next, Mendel crossed a Tt plant with itself. This led to four
genotypes, all equally likely: TT, Tt, tT and tt. Any genotype with the
dominant T allele results in a tall phenotype, while only the tt
genotype produces the short phenotype. When explained like that, the
3:1 ratio of tall to short discovered by Mendel makes perfect sense. It
also highlights the incredible leap of imagination Mendel made to
figure it all out.
Recessive traits

G enetic dominance is not integral to an allele; it is a phenomenon

that arises when two alleles meet. One allele may be dominant
over the other, in which case this second allele is termed ‘recessive’
to the dominant one. However, it is possible that the recessive allele
in this particular genotype is itself dominant over a third allele. For
example, the dark hair allele is dominant over the blonde allele, which
in turn is dominant over red hair.

Some traits, such as red hair, are entirely recessive but have little
effect on an organism’s chance of survival. Other examples, such as
albinism (opposite), can be more troublesome. In all cases, recessive
phenotypes are infrequent in the population because they require a
homozygous, double-recessive genotype. Many genotypes can contain
a recessive allele that remains hidden by a dominant partner. Only
when two heterozygous parents, or ‘carriers’, produce offspring is
there potential for the recessive phenotype to appear – seemingly out
of the blue.
Exploring the Variety of Random
Documents with Different Content
 § 30. Nomenclature by Position.
Two passages of Aristoxenus are quoted by Westphal in support of
his contention. The first (p. 6 Meib.) is one in which Aristoxenus
announces his intention to treat of Systems, their number and
nature: 'setting out their differences in respect of compass
(megethos), and for each compass the differences in form and
composition and position (tas te kata schêma kai kata synthesin
kai kata thesin), so that no element of melody,—either compass or
form or composition or position,—may be unexplained.' But the word
thesis, when applied to Systems, does not mean the 'position' of
single notes, but of groups of notes. Elsewhere (p. 54 Meib.) he
speaks of the position of tetrachords towards each other (tas tôn
tetrachordôn pros allêla theseis), laying it down that any two
tetrachords in the same System must be consonant either with each
other or with some third tetrachord. The other passage quoted by
Westphal (p. 69 Meib.) is also in the discussion of Systems.
Aristoxenus is pointing out the necessity of recognising that some
elements of melodious succession are fixed and limited, others are
unlimited:

kata men oun ta megethê tôn diastêmatôn kai tas tôn

phthongôn taseis apeira pôs phainetai einai ta peri
melos, kata de tas dynameis kai kata ta eidê kai kata tas
theseis peperasmena te kai tetagmena.
'In the size of the intervals and the pitch of the notes the
elements of melody seem to be infinite; but in respect of the
values (i.e. the relative places of the notes) and in respect of the
forms (i.e. the succession of the intervals) and in respect of the
positions they are limited and settled.'
Aristoxenus goes on to illustrate this by supposing that we wish to
continue a scale downwards from a pyknon or pair of small intervals
(Chromatic or Enharmonic). In this case, as the pyknon forms the
lower part of a tetrachord, there are two possibilities. If the next
lower tetrachord is disjunct, the next interval is a tone; if it is
conjunct, the next interval is the large interval of the genus (hê men
gar kata tonon eis diazeuxin agei to tou systêmatos eidos, hê
de kata thateron diastêma ho ti dêpot' echei megethos eis
synaphên). Thus the succession of intervals is determined by the
relative position of the two tetrachords, as to which there is a choice
between two definite alternatives. This then is evidently what is
meant by the words kata tas theseis [33]. On the other hand the
thesis of Ptolemy's nomenclature is the absolute pitch (Harm. ii. 5
pote men par' autên tên thesin, to oxyteron haplôs ê
baryteron, onomazomen), and this is one of the elements which
according to Aristoxenus are indefinite.
Westphal also finds the nomenclature by position implied in the
passage of the Aristotelian Problems (xix. 20) which deals with the
peculiar relation of the Mesê to the rest of the musical scale. The
passage has already been quoted and discussed (supra, p. 43), and it
has been pointed out that if the Mesê of the Perfect System (mesê
kata dynamin) is the key-note, the scale must have been an octave
of the a-species. If octaves of other species were used, as Westphal
maintains, it becomes necessary to take the Mesê of this passage to
be the mesê kata thesin, or Mesê by position. That is, Westphal is
obliged by his theory of the Modes to take the term Mesê in a sense
of which there is no other trace before the time of Ptolemy. But—
(1) It is highly improbable that the names of the notes—Mesê,
Hypatê, Nêtê and the rest—should have had two distinct meanings.
Such an ambiguity would have been intolerable, and only to be
compared with the similar ambiguity which Westphal's theory implies
in the use of the terms Dorian, &c.
(2) If the different species of the octave were the practically
important scales, as Westphal maintains, the position of the notes in
these scales must have been correspondingly important. Hence the
nomenclature by position must have been the more usual and
familiar one. Yet, as we have shown, it is not found in Aristotle,
Aristoxenus or Euclid—to say nothing of later writers.
(3) The nomenclature by position is an essential part of the scheme
of Keys proposed by Ptolemy. It bears the same relation to Ptolemy's
octaves as the nomenclature by 'value' bears to the old standard
octave and the Perfect System. It was probably therefore devised
about the time of Ptolemy, if not actually by him.
§ 31. Scales of the Lyre and Cithara.
The earliest evidence in practical music of any octaves other than
those of the standard System is to be found in the account given by
Ptolemy of certain scales employed on the lyre and cithara. According
to this account the scales of the lyre (the simpler and commoner
instrument) were of two kinds. One was Diatonic, of the 'colour' or
variety which Ptolemy recognises as the prevailing one, viz. the
'Middle Soft' or 'Tonic' (diatonon toniaion) [34]. The other was a
'mixture' of this Diatonic with the standard Chromatic (chrôma
suntonon): that is to say, the octave consisted of a tetrachord of
each genus. These octaves apparently might be of any species,
according to the key chosen [35]. On the cithara,—which was a more
elaborate form of lyre, confined in practice to professional musicians,
—six different octave scales were employed, each of a particular
species and key. They are enumerated and described by Ptolemy in
two passages (Harm. i. 16 and ii. 16), which in some points serve to
correct each other [36]. Of the six scales two are of the Hypo-dorian
or Common species (a-a). One of these, called tritai, is purely
Diatonic of the Middle Soft variety; the intervals expressed by
fractions are as follows:

a 9/8 b 28/27 c 8/7 d 9/8 e 28/27 f 8/7 g 9/8 a

The other, called tropoi or tropika, is a mixture, Middle Soft
Diatonic in the upper tetrachord, and Chromatic in the lower:

a 9/8 b 22/21 c 12/11 c♯ 7/6 e 28/27 f 8/7 g 9/8 a

Two scales are of the Dorian or e-species, viz. parypatai, a
combination of Soft and Middle Soft Diatonic:
 e 21/20 f 10/9 g 8/7 a 9/8 b 28/27 c 8/7 d 9/8 e
and lydia, in which the upper tetrachord is of the strict or 'highly
strung' Diatonic (diatonon syntonon—our 'natural' temperament):

e 28/27 f 8/7 g 9/8 a 9/8 b 16/15 c 9/8 d 10/9 e

Westphal (Harmonik und Melopöie, 1863, p. 255) supposes a
much deeper corruption. He would restore ta de lydia [kai
iastia hoi tou migmatos tou syntonou diatonou tou ... ta
de ...] hoi tou toniaiou diatonou tou Dôriou. This introduces
a serious discrepancy between the two passages, as the number
of scales in the second list is raised to eight (Westphal making
iastia and iastiaioliaia distinct scales, and furthermore
inserting a new scale, of unknown name). Moreover the
(unknown) scale of unmixed diatonon toniaion is out of its
place at the end of the list. Westphal's objection to lydia as the
name of a scale of the Dorian species of course only holds good
on his theory of the Modes.
The only other differences between the two passages are:
(1) In the scales of the lyre called malaka the admixture,
according to i. 16, is one of chrômatikon syntonon, according
to ii. 16 of chr. malakon. But, as Westphal shows, Soft
Chromatic is not admitted by Ptolemy as in practical use. It
would seem that in the second passage the copyist was led
astray by the word malaka just before.
(2) The iastia of i. 16 is called astiaioliaia in ii. 16. We need
not suppose the text to be faulty, since the two forms may have
been both in use.
Another point overlooked in Westphal's treatment is that
diatonon syntonon and d. ditoniaion are not really
distinguished by Ptolemy. In one passage (i. 16) he gives his
lydia and iastia as a mixture with d. syntonon, adding that in
practice it was d. ditoniaion. In the other (ii. 16) he speaks at
 once of d. ditoniaion. This consideration brings the two places
into such close agreement that any hypothesis involving
discrepancy is most improbable.

In practice it appears that musicians tuned the tetrachord b-e of this

scale with the Pythagorean two Major tones and leimma.
Of the remaining scales one, called hypertropa, is Phrygian in
species (d-d), and of the standard genus:

d 9/8 e 28/27 f 8/7 g 9/8 a 9/8 b 28/27 c 8/7 d

One, called iastia, or iastiaioliaia, is of the Hypo-phrygian or g-
species, the tetrachord b-e being 'highly strung' Diatonic or (in
practice) Pythagorean, viz.:

g 9/8 a 9/8 b 256/243 c 9/8 d 9/8 e 28/27 f 8/7 g

Regarding the tonality of these scales there is not very much to be
said. In the case of the Hypo-dorian and Dorian octaves it will be
generally thought probable that the key-note is a (the mesê kata
dynamin). If so, the difference between the two species is not one
of 'mode,'—in the modern sense,—but consists in the fact that in the
Hypo-dorian the compass of the melody is from the key-note
upwards, while in the Dorian it extends a Fourth below the key-note.
It is possible, however, that the lowest note (e) of the Dorian octave
was sometimes the key-note: in which case the mode was properly
Dorian. In the Phrygian octave of Ptolemy's description the key-note
cannot be the Fourth or Mesê kata thesin (g), since the interval g-c
is not consonant (9/8 × 9/8 × 28/27 being less than 4/3). Possibly
the lowest note (d) is the key-note; if so the scale is of the Phrygian
mode (in the modern sense). In the Hypo-phrygian octave there is a
similar objection to regarding the Mesê kata thesin (c) as the key-
note, and some probability in favour of the lowest note (g). If the
Pythagorean division of the tetrachord g-c were replaced by the
natural temperament, which the language used by Ptolemy [37] leads
us to regard as the true division, the scale would exhibit the intervals
—

g 5/4 b 6/5 d 7/6 f 8/7 g

which give the natural chord of the Seventh. This however is no more
than a hypothesis.
It evidently follows from all this that Ptolemy's octaves do not
constitute a system of modes. They are merely the groups of notes,
of the compass of an octave, which are most likely to be used in the
several keys, and which Ptolemy or some earlier theorist chose to call
by the names of those keys.
 § 32. Remains of Greek Music.
The extant specimens of Greek music are mostly of the second
century A.D., and therefore nearly contemporary with Ptolemy. The
most considerable are the melodies of three lyrical pieces or hymns,
viz. (1) a hymn to Calliope, (2) a hymn to Apollo (or Helios),—both
ascribed to a certain Dionysius,—and (3) a hymn to Nemesis,
ascribed to Mesomedes [38]. Besides these there are (4) some short
instrumental passages or exercises given by Bellermann's Anonymus
(pp. 94-96). And quite recently the list has been increased by (5) an
inscription discovered by Mr. W. M. Ramsay, which gives a musical
setting of four short gnomic sentences, and (6) a papyrus fragment
(now in the collection of the Arch-duke Rainer) of the music of a
chorus in the Orestes of Euripides. These two last additions to our
scanty stock of Greek music are set out and discussed by Dr. Wessely
of Vienna and M. Ruelle in the Revue des Études Grecques (V. 1892,
pp. 265-280), also by Dr. Otto Crusius in the Philologus, Vol. LII, pp.
160-200 [39].
The music of the three hymns is noted in the Lydian key (answering
to the modern scale with one ♭). The melody of the second hymn is
of the compass of an octave, the notes being those of the Perfect
System from Parhypatê Hypatôn to Tritê Diezeugmenôn (f-f with one
♭). The first employs the same octave with a lower note added, viz.
Hypatê Hypatôn (e): the third adds the next higher note, Paranêtê
Diezeugmenôn (g). Thus the Lydian key may be said, in the case of
the second hymn, and less exactly in the case of the two others, to
give the Lydian or c-species of the octave in the most convenient part
of the scale; just as on Ptolemy's system of Modes we should expect
it to do.
This octave, however, represents merely the compass (ambitus or
tessitura) of the melody: it has nothing to do with its tonality. In the
first two hymns, as Bellermann pointed out, the key-note is the
Hypatê Mesôn; and the mode—in the modern sense of that word—is
that of the octave e-e (the Dorian mode of Helmholtz's theory). In
the third hymn the key-note appears to be the Lichanos Mesôn, so
that the mode is that of g-g, viz. the Hypo-phrygian.
Of the instrumental passages given by the Anonymus three are
clearly in the Hypo-dorian or common mode, the Mesê (a) being the
key-note. (See Gevaert, i. p. 141.) A fourth (§ 104) also ends on the
Mesê, but the key-note appears to be the Parhypatê Mesôn (f).
Accordingly Westphal and Gevaert assign it to the Hypo-lydian
species (f-f). In Westphal's view the circumstance of the end of the
melody falling, not on the key-note, but on the Third or Mediant of
the octave, was characteristic of the Modes distinguished by the
prefix syntono-, and accordingly the passage in question is
pronounced by him to be Syntono-lydian. All those passages,
however, are mere fragments of two or three bars each, and are
quoted as examples of certain peculiarities of rhythm. They can
hardly be made to lend much support to any theory of the Modes.
The music of Mr. Ramsay's inscription labours under the same defect
of excessive shortness. If, however, we regard the four brief
sentences as set to a continuous melody, we obtain a passage
consisting of thirty-six notes in all, with a compass of less than an
octave, and ending on the lowest note of that compass. Unlike the
other extant specimens of Greek music it is written in the Ionian key
—a curious fact which has not been noticed by Dr. Wessely.

INSCRIPTION WITH MUSICAL NOTES.

 [Listen]

hos-on zês phai-nou.

mê-den hol-ôs sy ly-pou.
pros o-li-gon es-ti to zên.
to te-los ho chro-nos a-pai-tei.

The notes which enter into this melody form the scale f♯-g-a-b-c♯-d-
e-f ♯ , which is an octave of the Dorian species (e-e on the white
notes). Hence if f ♯, on which the melody ends, is the key-note, the
mode is the Dorian. On the other hand the predominant notes are
those of the triad a-c♯-e, which point to the key of a major, with the
difference that the Seventh is flat (g instead of g ♯). On this view the
music would be in the Hypo-phrygian mode.
However this may be, the most singular feature of this fragment
remains to be mentioned, viz. the agreement between the musical
notes and the accentuation of the words. We know from the
grammarians that an acute accent signified that the vowel was
sounded with a rise in the pitch of the voice, and that a circumflex
denoted a rise followed on the same syllable by a lower note—every
such rise and fall being quite independent both of syllabic quantity
and of stress or ictus. Thus in ordinary speech the accents formed a
species of melody,—logôdes ti melos, as it is called by Aristoxenus
[40]. When words were sung this 'spoken melody' was no longer
heard, being superseded by the melody proper. Dionysius of
Halicarnassus is at pains to explain (De Comp. Verb., c. 11), that the
melody to which words are set does not usually follow or resemble
the quasi-melody of the accents, e.g. in the following words of a
chorus in the Orestes of Euripides (ll. 140-142):—

siga siga leukon ichnos arbylês

tithete, mê ktypeite;
apoprobat' ekeis' apopro moi koitas,

he notices that the melody differs in several points from the spoken
accents: (1) the three first words are all on the same note, in spite of
the accents; (2) the last syllable of arbylês is as high as the second,
though that is the only accented syllable: (3) the first syllable of
tithete is lower than the two others, instead of being higher: (4) the
circumflex of ktypeite is lost (êphanistai), because the word is all
on the same pitch; (5) the fourth syllable of apoprobate is higher in
pitch, instead of the third. In Mr. Ramsay's inscription, however, the
music follows the accents as closely as possible. Every acute accent
coincides with a rise of pitch, except in hoson, which begins the
melody, and in esti, for which we should perhaps read the orthotone
esti. Of the four instances of the circumflex accent three exhibit the
two notes and the falling pitch which we expect. The interval is either
a major or a minor Third. In the other case (zês) the next note is a
Third lower: but it does not seem to belong to the circumflexed
syllable. All this cannot be accidental. It leads us to the conclusion
that the musical notes represent a kind of recitative, or imitation of
spoken words, rather than a melody in the proper sense of the term.
If any considerable specimen of the music of Euripides had survived,
it might have solved many of the problems with which we have been
dealing. The fragment before us extends over about six lines in
dochmiac metre (Orestes 338-343), with the vocal notation: but no
single line is entire. The key is the Lydian. The genus is either
Enharmonic or Chromatic. Assuming that it is Enharmonic—the
alternative adopted by Dr. Wessely—the characters which are still
legible may be represented in modern notation as follows:

[Listen]

(katolo)phy-ro-mai; ma-te-ros (haima sas ho d'

ana)bak-cheu-ei;
ho me-gas (olbos ou monimo)s en bro-tois;
a-na (de laiphos hôs ti)s a-ka-tou tho-as ti-
na(xas daimôn)
 kat-e-kly-sen (deinôn ponon) hôs pon-tou
labrois k.t.l.
It should be observed that in the fragment the line katolophyromai
katolophyromai comes before 338 (materos k.t.l.), not after it, as
in our texts [41].
The notes employed, according to the interpretation given above,
give the scale g-a-a*-a ♯ -d-e-e*. If the genus is Chromatic, as M.
Ruelle is disposed to think, they are g-a-a ♯ -b-d-e-f. When these
scales are compared with the Perfect System we find that they do not
entirely agree with it. Whether the genus is Enharmonic or Chromatic
the notes from a to e* (or f) answer to those of the Perfect System
(of the same genus) from Hypatê Mesôn to Tritê Diezeugmenôn. But
in either case the lowest note (g) finds no place in the System, since
it can only be the Diatonic Lichanos Hypatôn. It is possible, however,
that the scale belongs to the period when the original octave had
been extended by the addition of a tone below the Hypatê—the note,
in fact, which we have already met with under the name of Hyper-
hypatê (p. 39). Thus the complete scale may have consisted of the
disjunct tetrachords a-d and e-a, with the tone g-a. It may be
observed here that although the scale in question does not fit into
the Perfect System, it conforms to the general rules laid down by
Aristoxenus for the melodious succession of intervals. It is
unnecessary therefore to suppose (as Dr. Wessely and M. Ruelle do)
that the scale exhibits a mixture of different genera.
It must be vain to attempt to discover the tonality of a short
fragment which has neither beginning nor end. The only group of
notes which has the character of a cadence is that on the word
(olo)phypomai, and again on the words en brotois, viz. the notes
a ♯ a* a (if the genus is the Enharmonic). The same notes occur in
reversed order on akatou and (kat)eklusen. This seems to bear
out the common view of the Enharmonic as produced by the
introduction of an 'accidental' or passing note. It will be seen, in fact,
that the Enharmonic notes (a* and e*) only occur before or after the
'standing' notes (a and e).
Relying on the fact that the lowest note is g, Dr. Wessely and M.
Ruelle pronounce the mode to be the Phrygian (g-g in the key with
one ♭ , or d-d in the natural key). I have already put forward a
different explanation of this g, and will only add here that it occurs
twice in the fragment, both times on a short syllable [42]. The
important notes, so far as the evidence goes, are a, which twice
comes at the end of a verse (with a pause in the sense), and e,
which once has that position. If a is the key-note, the mode—in the
modern sense—is Dorian (the e-species). If e is the key-note, it is
Mixo-lydian (the b-species).
 § 33. Modes of Aristides
Quintilianus.
The most direct testimony in support of the view that the ancient
Modes were differentiated by the succession of their intervals has still
to be considered. It is the account given by Aristides Quintilianus (p.
21 Meib.) of the six Modes (harmoniai) of Plato's Republic. After
describing the genera and their varieties the 'colours,' he goes on to
say that there were other divisions of the tetrachord (tetrachordikai
diaireseis) which the most ancient musicians used for the
harmoniai, and that these were sometimes greater in compass than
the octave, sometimes less. He then gives the intervals of the scale
for each of the six Modes mentioned by Plato, and adds the scales in
the ancient notation. They are of the Enharmonic genus, and may be
represented by modern notes as follows:—
Mixo-lydian b-b*-c-d-e-e*-f-b
Syntono-lydian e-e*-f-a-c
Phrygian d-e-e*-f-a-b-b*-c-d
Dorian d-e-e*-f-a-b-b*-c-e
Lydian e*-f-a-b-b*-c-e-e*
Ionian e-e*-f-a-c-d
Comparing these scales with the Species of the Octave, we find a
certain amount of correspondence. As has been already noticed (p.
22), the names Syntono-lydian and Lydian answer to the ordinary
Lydian and Hypo-lydian respectively. Accordingly the Lydian of
Aristides agrees with the Hypo-lydian species as given in the pseudo-
Euclidean Introductio. The Dorian of Aristides is the Dorian species of
the Introductio, but with an additional note, a tone below the Hypatê.
The Phrygian of Aristides is not the Enharmonic Phrygian species; but
it is derived from the diatonic Phrygian octave d-e-f-g-a-b-c-d by
inserting the enharmonic notes e* and b*, and omitting the diatonic
g. By a similar process the Mixo-lydian of Aristides may be derived
from the diatonic octave b-b, except that a as well as g is omitted,
and on the other hand d is retained. If the scale of the Syntono-
lydian is completed by the lower c (as analogy would require), it will
answer similarly to the Lydian species (c-c).
 § 34. Credibility of Aristides
Quintilianus.
But what weight can be given to Aristides as an authority on the
music of the time of Plato? The answer to this question depends
upon several considerations.
1. The date of Aristides is unknown. He is certainly later than Cicero,
since he quotes the De Republica (p. 70 Meib.). From the
circumstance that he makes no reference to the musical innovations
of Ptolemy it has been supposed that he was earlier than that writer.
But, as Aristides usually confines himself to the theory of Aristoxenus
and his school, the argument from silence is not of much value. On
the other hand he gives a scheme of notation containing two
characters, and , which extend the scale two successive semi-
tones beyond the lowest point of the notation given by Alypius [43].
For this reason it is probable that Aristides is one of the latest of the
writers on ancient music.
2. The manner in which Aristides introduces his information about the
Platonic Modes is highly suspicious. He has been describing the
various divisions of the tetrachord according to the theory of
Aristoxenus, and adds that there were anciently other divisions in
use. So far Aristides is doubtless right, since Aristoxenus himself says
that the divisions of the tetrachord are theoretically infinite in number
(p. 26 Meib.),—that it is possible, for example, to combine the
Parhypatê of the Soft Chromatic with the Lichanos of the Diatonic (p.
52 Meib.). But all this concerns the genus of the scale, and has
nothing to do with the species of the Octave, with which Aristides
proceeds to connect it. It follows either that there is some confusion
in the text, or that Aristides was compiling from sources which he did
not understand.
3. The Platonic Modes were a subject of interest to the early musical
writers, and were discussed by Aristoxenus himself (Plut. de Mus. c.
17). If Aristoxenus had had access to such an account as we have in
Aristides, we must have found some trace of it, either in the extant
Harmonics or in the quotations of Plutarch and other compilers.
4. Of the four scales which extend to the compass of an octave, only
one, viz. the Dorian, conforms to the rules which are said by
Aristoxenus to have prevailed in early Greek music. The Phrygian
divides the Fourth a-d into four intervals instead of three, by the
sequence a b b* c d. As has been observed, it is neither the
Enharmonic Phrygian species (c e e* f a b b* c), nor the Diatonic d-d,
but a mixture of the two. Similarly the Mixo-lydian divides the Fourth
b-e into four intervals (b b* c d e), by introducing the purely Diatonic
note d. The Lydian is certainly the Lydian Enharmonic species of the
pseudo-Euclid; but we can hardly suppose that it existed in practical
music. Aristoxenus lays it down emphatically that a quarter-tone is
always followed by another: and we cannot imagine a scale in which
the highest and lowest notes are in no harmonic relation to the rest.
5. Two of the scales are incomplete, viz. the Ionian, which has six
notes and the compass of a Seventh, and the Syntono-lydian, which
consists of five notes, with the compass of a Minor Sixth. We
naturally look for parallels among the defective scales noticed in the
Problems and in Plutarch's dialogues. But we find little that even
illustrates the modes of Aristides. The scales noticed in the Problems
(xix. 7, 32, 47) are hepta-chord, and generally of the compass of an
octave. In one passage of Plutarch (De Mus. c. 11) there is a
description—quoted from Aristoxenus—of an older kind of
Enharmonic, in which the semitones had not yet been divided into
quarter-tones. In another chapter (c. 19) he speaks of the omission
of the Tritê and also of the Nêtê as characteristic of a form of music
called the spondeiakos tropos. It may be said that in the Ionian
and Syntono-lydian of Aristides the Enharmonic Tritê (b*) and the
Nêtê (e) are wanting. But the Paramesê (b) is also wanting in both
these modes. And the Ionian is open to the observation already made
with regard to the Phrygian, viz. that the two highest notes (c d)
involve a mixture of Diatonic with Enharmonic scale. We may add
that Plutarch (who evidently wrote with Aristoxenus before him) gives
no hint that the omission of these notes was characteristic of any
particular modes.
6. It is impossible to decide the question of the modes of Aristides
without some reference to another statement of the same author. In
the chapter which treats of Intervals (pp. 13-15 Meib.) he gives the
ancient division of two octaves, the first into dieses or quarter-tones,
the second into semitones. The former of these (hê para tois
archaiois kata dieseis harmonia) is as follows:

After every allowance has been made for the probability that these
signs or some of them have reached us in a corrupt form, it is
impossible to reduce them to the ordinary notation, as Meibomius
sought to do. The scholar who first published them as they stand in
the MSS. (F. L. Perne, see Bellermann, Tonleitern, p. 62) regarded
them as a relic of a much older system of notation. This is in
accordance with the language of Aristides, and indeed is the only
view consistent with a belief in their genuineness. They are too like
the ordinary notation to be quite independent, and cannot have been
put forward as an improvement upon it. Are they, then, earlier?
Bellermann has called our attention to a peculiarity which seems fatal
to any such claim. They consist, like the ordinary signs, of two sets,
one written above the other, and in every instance one of the pair is
simply a reversed or inverted form of the other. With the ordinary
signs this is not generally the case, since the two sets, the vocal and
instrumental notes, are originally independent. But it is the case with
the three lowest notes, viz. those which were added to the series at a
later time. When these additional signs were invented the vocal and
instrumental notes had come to be employed together. The inventor
therefore devised a pair of signs in each case, and not unnaturally
made them correspond in form. In the scale given by Aristides this
correspondence runs through the whole series, which must therefore
be of later date. But if this is so, the characters can hardly represent
a genuine system of notation. In other words, Aristides must have
been imposed upon by a species of forgery.
7. Does the fragment of the Orestes tell for or against the Modes
described by Aristides?
The scale which is formed by the notes of the fragment agrees, so far
as it extends, with two of the scales now in question, viz. the
Phrygian and the Dorian. Taking the view of its tonality expressed in
the last chapter (p. 93), we should describe it as the Dorian scale of
Aristides with the two highest notes omitted. The omission, in so
short a fragment, is of little weight; and the agreement in the use of
an additional lower note (Hyper-hypatê) is certainly worth notice. On
the other hand, the Dorian is precisely the mode, of those given in
the list of Aristides, which least needs defence, as it is the most
faithful copy of the Perfect System. Hence the fact that it is verified
by an actual piece of music does not go far in support of the other
scales in the same list.
If our suspicions are well-founded, it is evident that they seriously
affect the genuineness of all the antiquarian learning which Aristides
sets before his readers, and in particular of his account of the
Platonic modes. I venture to think that they go far to deprive that
account of the value which it has been supposed to have for the
history of the earliest Greek music.
For the later period, however, to which Aristides himself belongs,
these apocryphal scales are a document of some importance. The
fact that they do not agree entirely with the species of the Octave as
given by the pseudo-Euclid leads us to think that they may be
influenced by scales used in actual music. This applies especially to
the Phrygian, which (as has been shown) is really diatonic. The
Ionian, again, is perhaps merely an imperfect form of the same scale,
viz. the octave d-d with lower d omitted. And the Syntono-lydian may
be the Lydian diatonic octave c-c with a similar omission of the lower
c.
§ 35. Evidence for Scales of different
species.
The object of the foregoing discussion has been to show, in the first
place, that there was no such distinction in ancient Greek music as
that which scholars have drawn between Modes (harmoniai) and
Keys (tonoi or tropoi): and, in the second place, that the musical
scales denoted by these terms were primarily distinguished by
difference of pitch,—that in fact they were so many keys of the
standard scale known in its final form as the Perfect System. The
evidence now brought forward in support of these two propositions is
surely as complete as that which has been allowed to determine any
question of ancient learning.
It does not, however, follow that the Greeks knew of no musical
forms analogous to our Major and Minor modes, or to the mediaeval
Tones. On the contrary, the course of the discussion has led us to
recognise distinctions of this kind in more than one instance. The
doctrine against which the argument has been mainly directed is not
that ancient scales were of more than one species or 'mode' (as it is
now called), but that difference of species was the basis of the
ancient Greek Modes. This will become clear if we bring together all
the indications which we have observed of scales differing from each
other in species, that is, in the order of the intervals in the octave. In
doing so it will be especially important to be guided by the principle
which we laid down at the outset, of arranging our materials
according to chronology, and judging of each piece of evidence
strictly with reference to the period to which it belongs. It is only thus
that we can hope to gain a conception of Greek music as the living
and changing thing that we know it must have been.
1. The principal scale of Greek music is undoubtedly of the Hypo-
dorian or common species. This is sufficiently proved by the facts (1)
that two octaves of this species (a-a) constitute the scale known as
the Greater Perfect System, and (2) that the central a of this system,
called the Mesê, is said to have been the key-note, or at least to have
had the kind of importance in the scale which we connect with the
key-note (Arist. Probl. xix. 20). This mode, it is obvious, is based on
the scale which is the descending scale of the modern Minor mode. It
may therefore be identified with the Minor, except that it does not
admit the leading note.
It should be observed that this mode is to be recognised not merely
in the Perfect System but equally in the primitive octave, of the form
e-e, out of which the Perfect System grew. The important point is the
tonic character of the Mesê (a), and this, as it happens, rests upon
the testimony of an author who knows the primitive octave only. The
fact that that octave is of the so-called Dorian species does not alter
the mode (as we are now using that term), but only the compass of
the notes employed.
The Hypo-dorian octave is seen in two of the scales of the cithara
given by Ptolemy (p. 85), viz. those called tritai and tropoi, and the
Dorian octave (e-e) in two scales, parupatai and ludia. It is very
possible (as was observed in commenting on them) that the two
latter scales were in the key of a, and therefore Hypo-dorian in
respect of mode. The Hypo-dorian mode is also exemplified by three
at least of the instrumental passages given by the Anonymus (supra,
p. 89).
2. The earliest trace of a difference of species appears to be found in
the passage on the subject of the Mixo-lydian mode quoted above (p.
24) from Plutarch's Dialogue on Music. In that mode, according to
Plutarch, it was discovered by a certain Lamprocles of Athens that the
Disjunctive Tone was the highest interval, that is to say, that the
octave in reality consisted of two conjunct tetrachords and a tone:
 [Listen]

As the note which is the meeting-point of the two tetrachords is

doubtless the key-note, we shall not be wrong in making it the Mesê,
and thus finding the octave in question in the Perfect System and in
the oldest part of it, viz. the tetrachords Mesôn and Synêmmenôn,
with the Nêtê Diezeugmenôn. How then did this octave come to be
recognised by Lamprocles as distinctively Mixo-lydian? We cannot tell
with certainty, because we do not know what the Mixo-lydian scale
was before his treatment of it. Probably, however, the answer is to be
sought in the relation in respect of pitch between the Dorian and
Mixo-lydian keys. These, as we have seen (p. 23), were the keys
chiefly employed in tragedy, and the Mixo-lydian was a Fourth higher
than the other. Now when a scale consisting of white notes is
transposed to a key a Fourth higher, it becomes a scale with one ♭. In
ancient language, the tetrachord Synêmmenôn (a-b ♭ -c-d) takes the
place of the tetrachord Diezeugmenôn. In some such way as this the
octave of this form may have come to be associated in a special way
with the use of the Mixo-lydian key.
However this may be, the change from the tetrachord Diezeugmenôn
to the tetrachord Synêmmenôn, or the reverse, is a change of mode
in the modern sense, for it is what the ancients classified as a change
of System (metabolê kata systêma) [44]. Nor is it hard to
determine the two 'modes' concerned, if we may trust to the
authority of the Aristotelian Problems (l. c.) and regard the Mesê as
always the key-note. For if a is kept as the key-note, the octave a-a
with one ♭ is the so-called Dorian (e-e on the white notes). In this
way we arrive at the somewhat confusing result that the ancient
Dorian species (e-e but with a as key-note) yields the Hypo-dorian or
modern Minor mode: while the Dorian mode of modern scientific
theory [45] has its ancient prototype in the Mixo-lydian species, viz.
the octave first brought to light by Lamprocles. The difficulty of
course arises from the species of the Octave being classified
according to their compass, without reference to the tonic character
of the Mesê.
The Dorian mode is amply represented in the extant remains of
Greek music. It is the mode of the two compositions of Dionysius, the
Hymn to Calliope and the Hymn to Apollo (p. 88), perhaps also of Mr.
Ramsay's musical inscription (p. 90). It would have been satisfactory
if we could have found it in the much more important fragment of the
Orestes. Such indications as that fragment presents seem to me to
point to the Dorian mode (Mixo-lydian of Lamprocles).
3. The scales of the cithara furnish one example of the Phrygian
species (d-d), and one of the Hypo-phrygian (g-g): but we have no
means of determining which note of the scale is to be treated as the
key-note.
In the Hymn to Nemesis, however, in spite of the incomplete form in
which it has reached us, there is a sufficiently clear example of the
Hypo-phrygian mode. It has been suggested as possible that the
melody of Mr. Ramsay's inscription is also Hypo-phrygian, and if so
the evidence for the mode would be carried back to the first century.
The Hypo-phrygian is the nearest approach made by any specimen of
Greek music to the modern Major mode,—the Lydian or c-species not
being found even among the scales of the cithara as given by
Ptolemy. It is therefore of peculiar interest for musical history, and we
look with eagerness for any indication which would allow us to
connect it with the classical period of Greek art. One or two sayings
of Aristotle have been thought to bear upon this issue.
The most interesting is a passage in the Politics (iv. 3, cp. p. 13),
where Aristotle is speaking of the multiplicity of forms of government,
and showing how a great number of varieties may nevertheless be
brought under a few classes or types. He illustrates the point from
the musical Modes, observing that all constitutions may be regarded
as either oligarchical (government by a minority) or democratical
(government by the majority), just as in the opinion of some
musicians (hôs phasi tines) all modes are essentially either Dorian
or Phrygian. What, then, is the basis of this grouping of certain
modes together as Dorian, while the rest are Phrygian in character?
According to Westphal it is a form of the opposition between the true
Hellenic music, represented by Dorian, and the foreign music, the
Phrygian and Lydian, with their varieties. Moreover, it is in his view
virtually the same distinction as that which obtains in modern music
between the Minor and the Major scales [46]. This account of the
matter, however, is not supported by the context of the passage.
Aristotle draws out the comparison between forms of government
and musical modes in such a way as to make it plain that in the case
of the modes the distinction was one of pitch (tas suntonôteras ...
tas d' aneimenas kai malakas). The Dorian was the best, because
the highest, of the lower keys,—the others being Hypo-dorian (in the
earlier sense, immediately below Dorian), and Hypo-phrygian—while
Phrygian was the first of the higher series which took in Lydian and
Mixo-lydian. The division would be aided, or may even have been
suggested, by the circumstance that it nearly coincided with the
favourite contrast of Hellenic and 'barbarous' modes [47]. There is
another passage, however, which can hardly be reconciled with a
classification according to pitch alone. In the chapters dealing with
the ethical character of music Aristotle dwells (as will be
remembered) upon the exciting and orgiastic character of the
Phrygian mode, and notices its especial fitness for the dithyramb.
This fitness or affinity, he says, was so marked that a poet who tried
to compose a dithyramb in another mode found himself passing
unawares into the Phrygian (Pol. viii. 7). It is natural to understand
this of the use of certain sequences of intervals, or of cadences, such
as are characteristic of a 'mode' in the modern sense of the word,
rather than of a change of key. If this is so we may venture the
further hypothesis that the Phrygian music, in some at least of its
forms, was distinguished not only by pitch, but also by the more or
less conscious use of scales which differed in type from the scale of
the Greek standard system.
It may be urged that this hypothesis is inconsistent with our
interpretation of the passage of the Problems about the tonic
character of the Mesê. If a is key-note, it was argued, the mode is
that of the a-species (Hypo-dorian, our Minor), or at most—by
admitting the tetrachord Synêmmenôn—it includes the e-species
(Dorian of Helmholtz). The answer may be that the statement of the
Problems is not of this absolute kind. It is not the statement of a
technical writer, laying down definite rules, but is a general
observation, or at best a canon of taste. We are not told how the
predominance of the Mesê is shown in the form of the melody.
Moreover this predominance is not said to be exercised in music
generally, but in all good music (panta gar ta chrêsta melê
pollakis tê mesê chrêtai). This may mean either that tonality in
Greek music was of an imperfect kind, a question of style and taste
rather than of fixed rule, or that they occasionally employed modes of
a less approved stamp, unrecognised in the earlier musical theory.
 § 36. Conclusion.
The considerations set forth in the last chapter seem to show that if
difference of mode or species cannot be entirely denied of the
classical period of Greek music, it occupied a subordinate and almost
unrecognised place.
The main elements of the art were, (1) difference of genus,—the
sub-divisions of the tetrachord which Aristoxenus and Ptolemy alike
recognise, though with important discrepancies in detail; (2)
difference of pitch or key; and (3) rhythm. Passing over the last, as
not belonging to the subject of Harmonics, we may now say that
genus and key are the only grounds of distinction which are evidently
of practical importance. No others were associated with the early
history of the art, with particular composers or periods, with
particular instruments, or with the ethos of music. This, however, is
only true in the fullest sense of Greek music before the time of
Ptolemy. The main object of Ptolemy's reform of the keys was to
provide a new set of scales, each characterised by a particular
succession of intervals, while the pitch was left to take care of itself.
And it is clear, especially from the specimens which Ptolemy gives of
the scales in use in his time, that he was only endeavouring to
systematise what already existed, and bring theory into harmony with
the developments of practice. We must suppose, therefore, that the
musical feeling which sought variety in differences of key came to
have less influence on the practical art, and that musicians began to
discover, or to appreciate more than they had done, the use of
different 'modes' or forms of the octave scale. Along with this change
we have to note the comparative disuse of the Enharmonic and
Chromatic divisions of the tetrachord. The Enharmonic, according to
Ptolemy, had ceased to be employed. Of the three varieties of
Chromatic given by Aristoxenus only one remains on Ptolemy's list,
and that the one which in the scheme of Aristoxenus involved no
interval less than a semitone. And although Ptolemy distinguished at
least three varieties of Diatonic, it is worth notice that only one of
these was admitted in the tuning of the lyre,—the others being
confined to the more elaborate cithara. In Ptolemy's time, therefore,
music was rapidly approaching the stage in which all its forms are
based upon a single scale—the natural diatonic scale of modern
Europe.
In the light of these facts it must occur to us that Westphal's theory
of seven modes or species of the Octave is really open to an a priori
objection as decisive in its nature as any of the testimony which has
been brought against it. Is it possible, we may ask, that a system of
modes analogous to the ecclesiastical Tones can have subsisted along
with a system of scales such as the genera and 'colours' of early
Greek music? The reply may be that Ptolemy himself combines the
two systems. He supposes five divisions of the tetrachord, and seven
modes based upon so many species of the Octave—in all thirty-five
different scales (or seventy, if we bring in the distinction of octaves
apo nêtês and apo mesês). But when we come to the scales
actually used on the chief Greek instrument, the cithara, the number
falls at once to six. Evidently the others, or most of them, only
existed on paper, as the mathematical results of certain assumptions
which Ptolemy had made. And if this can be said of Ptolemy's theory,
what would be the value of a similar scheme combining the modes
with the Enharmonic and the different varieties of the Chromatic
genus? The truth is, surely, that such a scheme tries to unite
elements which belong to different times, which in fact are the
fundamental ideas of different stages of art.
The most striking characteristic of Greek music, especially in its
earlier periods, is the multiplicity and delicacy of the intervals into
which the scale was divided. A sort of frame-work was formed by the
division of the octave into tetrachords, completed by the so-called
disjunctive tone; and so far all Greek music was alike. But within the
tetrachord the reign of diversity was unchecked. Not only were there
recognised divisions containing intervals of a fourth, a third, and even
three-eighths of a tone, but we gather from several things said by
Aristoxenus that the number of possible divisions was regarded as
theoretically unlimited. Thus he tells us that there was a constant
tendency to flatten the 'moveable' notes of the Chromatic genus, and
thus diminish the small intervals, for the sake of 'sweetness' or in
order to obtain a plaintive tone [48];— that the Lichanos of a
tetrachord may in theory be any note between the Enharmonic
Lichanos (f in the scale e-e*-f-a) and the Diatonic (g in the scale e-f-
g-a) [49];— and that the magnitude of the smaller intervals and
division of the tetrachord generally belongs to the indefinite or
indeterminate element in music [50]. Moreover, in spite of the disuse
of several of the older scales, much of this holds good for the time of
Ptolemy. The modern diatonic scale is fully recognised by him, but
only as one of several different divisions. And the division which he
treats as the ordinary or standard form of the octave is not the
modern diatonic scale, but one of the so-called 'soft' or flattened
varieties. It is clear that in the best periods of Greek music these
refinements of melody, which modern musicians find scarcely
conceivable, were far from being accidental or subordinate features.
Rather, they were as much bound up with the fundamental nature of
that music as complex harmony is with the music of modern Europe.
The mediaeval modes or Tones, on the other hand, are essentially
based on the diatonic scale,—the scale that knows only of tones and
semitones. To suppose that they held in the earliest Greek music the
prominent place which we find assigned to the ancient Modes or
harmoniai is to suppose that the art of music was developed in
Greece in two different directions, under the influence of different
and almost opposite ideas. Yet nothing is more remarkable in all
departments of Greek art than the strictness with which it confines
itself within the limits given once for all in the leading types, and the
consequent harmony and consistency of all the forms which it takes
in the course of its growth.
The dependence of artistic forms in their manifold developments
upon a central governing idea or principle has never been more
luminously stated than by the illustrious physicist Helmholtz, in the
thirteenth chapter of his Tonempfindungen. I venture to think that in
applying that truth to the facts of Greek music he was materially
hindered by the accepted theory of the Greek modes. The scales
which he analyses under that name were certainly the basis of all
music in the Middle Ages, and are much more intelligible as such
than in relation to the primitive Greek forms of the art [51].
 § 37. Epilogue—Speech and Song.
Several indications combine to make it probable that singing and
speaking were not so widely separated from each other in Greek as in
the modern languages with which we are most familiar.
(1) The teaching of the grammarians on the subject of accent points
to this conclusion. Our habit of using Latin translations of the terms
of Greek grammar has tended to obscure the fact that they belong in
almost every case to the ordinary vocabulary of music. The word for
'accent' (tonos) is simply the musical term for 'pitch' or 'key.' The
words 'acute' (oxys) and 'grave' (barys) mean nothing more than
'high' and 'low' in pitch. A syllable may have two accents, just as in
music a syllable may be sung with more than one note. Similarly the
'quantity' of each syllable answers to the time of a musical note, and
the rule that a long syllable is equal to two short ones is no doubt
approximately correct. Consequently every Greek word (enclitics
being reckoned as parts of a word) is a sort of musical phrase, and
every sentence is a more or less definite melody—logôdes ti melos,
as it is called by Aristoxenus (p. 18 Meib.). Moreover the accent in the
modern sense, the ictus or stress of the voice, appears to be quite
independent of the pitch or 'tonic' accent: for in Greek poetry the
ictus (arsis) is determined by the metre, with which the tonic accent
evidently has nothing to do. In singing, accordingly, the tonic accents
disappear; for the melody takes their place, and gives each syllable a
new pitch, on which (as we shall presently see) the spoken pitch has
no influence. The rise and fall of the voice in ordinary speaking is
perceptible enough in English, though it is more marked in other
European languages. Helmholtz tells us—with tacit reference to the
speech of North Germany—that an affirmative sentence generally
ends with a drop in the tone of about a Fourth, while an interrogative
is marked by a rise which is often as much as a Fifth [52]. In Italian
the interrogative form is regularly given, not by a particle or a change
in the order of the words, but by a rise of pitch. The Gregorian
church music, according to a series of rules quoted by Helmholtz (l.
c.), marked a comma by a rise of a Tone, a colon by a fall of a
Semitone; a full stop by a Tone above, followed by a Fourth below,
the 'reciting note'; and an interrogation by a phrase of the form d b c
d (c being the reciting note).
These examples, however, do little towards enabling modern scholars
to form a notion of the Greek system of accentuation. In these and
similar cases it is the sentence as a whole which is modified by the
tonic accent, whereas in Greek it is the individual word. It is true that
the accent of a word may be affected by its place in the sentence: as
is seen in the loss of the accent of oxytone words when not followed
by a pause, in the anastrophe of prepositions, and in the treatment of
the different classes of enclitics. But in all these instances it is the
intonation of the word as such, not of the sentence, which is
primarily concerned. What they really prove is that the musical accent
is not so invariable as the stress accent in English or German, but
may depend upon the collocation of the word, or upon the degree of
emphasis which it has in a particular use.
(2) The same conclusion may be drawn from the terms in which the
ancient writers on music endeavour to distinguish musical and
ordinary utterance.
Aristoxenus begins his Harmonics by observing that there are two
movements of the voice, not properly discriminated by any previous
writer; namely, the continuous, which is the movement characteristic
of speaking, and the discrete or that which proceeds by intervals, the
movement of singing. In the latter the voice remains for a certain
time on one note, and then passes by a definite interval to another.
In the former it is continually gliding by imperceptible degrees from
higher to lower or the reverse [53]. In this kind of movement the rise
and fall of the voice is marked by the accents (prosôdiai), which
accordingly form the melody, as it may be called, of spoken utterance
[54]. Later writers state the distinction in much the same language.
Nicomachus tells us that the two movements were first discriminated
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