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BY THE PEOPLE
DEBATING AMERICAN GOVERNMENT
BY THE PEOPLE
DEBATING AMERICAN GOVERNMENT | BRIEF
FOURTH EDITION

JAMES A. MORONE
Brown University
ROGAN KERSH
Wake Forest University
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address above.

You must not circulate this work in any other form and you must impose this same condition
on any acquirer.

Library of Congress Cataloging-in-Publication Data

Names: Morone, James A., 1951- author. | Kersh, Rogan, author.


Title: By the people : debating American government / James A. Morone, Brown University
Rogan Kersh, Wake Forest University.
Description: Brief Fourth Edition. | New York : Oxford University Press, [2018] | Include
bibliographical references and index.
Identifiers: LCCN 2018049759| ISBN 9780190928728 (Paperback) |
ISBN 9780190928759 (Looseleaf) | ISBN 9780190928735 (ebook) | Subjects: LCSH: United
States. Constitution. | United States—Politics and government—Textbooks.
Classification: LCC JK276 .M67 2019 | DDC 320.473—dc23
LC record available at https://lccn.loc.gov/2018049759

987654321
Printed by LSC Communications, Inc., United States of America
Many teachers and colleagues inspired us. We dedicate this book to
four who changed our lives. Their passion for learning and teaching set
the standard we aim for every day—and on every page that follows.

Richard O’Donnell
Murray Dry
Jim Barefield
Rogers Smith
By the People comes from the Gettysburg Address. Standing on the
battlefield at Gettysburg, President Abraham Lincoln delivered wha
may be the most memorable presidential address in American history—
defining American government as a government “of the people, by the
people, for the people.” Here is the full address.

F our score and seven years ago our fathers brought forth on this
continent, a new nation, conceived in liberty, and dedicated to the
proposition that all men are created equal.

Now we are engaged in a great civil war, testing whether that nation
or any nation so conceived and so dedicated, can long endure. We are
met on a great battle-field of that war. We have come to dedicate a
portion of that field, as a final resting place for those who here gave
their lives that that nation might live. It is altogether fitting and prope
that we should do this.

But, in a larger sense, we can not dedicate—we can not consecrate—


we can not hallow—this ground. The brave men, living and dead, who
struggled here, have consecrated it, far above our poor power to add
or detract. The world will little note, nor long remember what we say
here, but it can never forget what they did here. It is for us the living
rather, to be dedicated here to the unfinished work which they who
fought here have thus far so nobly advanced. It is rather for us to be
here dedicated to the great task remaining before us—that from these
honored dead we take increased devotion to that cause for which they
gave the last full measure of devotion—that we here highly resolve
that these dead shall not have died in vain—that this nation, unde
God, shall have a new birth of freedom—and that government of the
people, by the people, for the people, shall not perish from the earth.
Brief Contents

About the Authors


Preface
Acknowledgments

PART I IDEAS AND RIGHTS

1 Ideas That Shape American Politics

2 The Constitution

3 Federalism and Nationalism

4 Civil Liberties

5 The Struggle for Civil Rights

PART II POLITICAL BEHAVIOR

6 Public Opinion and Political Participation

7 Media, Technology, and Government

8 Campaigns and Elections

9 Interest Groups and Political Parties


PART III POLITICAL INSTITUTIONS

1 Congress
0
1 The Presidency
1
1 Bureaucracy
2
1 The Judicial Branch
3
PART IV POLICYMAKING

1 Domestic and Foreign Policy


4

APPENDIX I
The Declaration of Independence

APPENDIX II
The Constitution of the United States of America

APPENDIX III
The Federalist Papers nos. 1, 10, and 51

Glossary
Notes
Credits
Index
Presidential Elections, Congressional Control, 1789–2019
Contents

About the Authors


Preface
Acknowledgments

PART I IDEAS AND RIGHTS

1 Ideas That Shape American Politics


BY THE NUMBERS American Ideas

The Spirit of American Politics


Who Governs?
How Does American Politics Work?
Ideas
Institutions
Interests
Individuals
What Does Government Do?
Who Are We?

A Nation of Ideas
Liberty
“The Land of the Free”
The Two Sides of Liberty

NEGATIVE VERSUS
WHAT DO YOU THINK?
POSITIVE LIBERTY
The Idea of Freedom Is Always Changing

Self-Rule
One Side of Self-Rule: Democracy
Another Side of Self-Rule: A Republic
A Mixed System

Limited Government
The Origins of Limited Government
And Yet . . . the United States Has a Big Government
Limits on Government Action
When Ideas Clash: Self-Rule and Limited Government

WHAT DO YOU THINK? SELF-RULE VERSUS LIMITED


GOVERNMENT
Individualism
Community Versus Individualism
The Roots of American Individualism: Opportunity and Discord
Golden Opportunity
Social Conflict
Who We Are: Individualism and Solidarity?

WHAT DO YOU THINK? INDIVIDUALISM VERSUS


SOLIDARITY
The American Dream
Spreading the Dream
Challenging the Dream
Is the System Tilted Toward the Wealthy?

INFO DATAMost Americans Believe There Is:


“Opportunity to Get Ahead”
Does the American Dream Promote the Wrong Values?

Equality
Three Types of Equality
How Much Economic Inequality Is Too Much?
Opportunity or Outcome?

Religion
Still a Religious Country
So Many Religions
The Politics of Religion

How Do Ideas Affect Politics?


Ideas in American Culture
The Ideas in Political Institutions
Culture or Institutions?

Conclusion: Culture and Institutions, Together


Chapter Summary
Key Terms
Study Questions

2 The Constitution
The Colonial Roots of the Constitution
BY THE NUMBERS The Constitution

Why the Colonists Revolted


The Colonial Complaint: Representation
The Conflict Begins with Blood on the Frontier
The Stamp Tax and the First Hints of Independence
The Townshend Acts Worsen the Conflict
The Boston Tea Party
Revolution!
A Long Legacy

The Declaration of Independence


The Principle: “We Hold These Truths . . .”
Grievances

The First American Government: The Articles of


Confederation
Independent States
The National Government
Some Success . . .
. . . And Some Problems

WHAT DO YOU THINK? YOUR ADVICE IS NEEDED


Secrecy

The Constitutional Convention


How Much Power to the People?
National Government Versus State Government
Big States Versus Small States
The Virginia Plan
The New Jersey Plan
The Connecticut Compromise
The President
Committee or Individual?
The Electoral College
Separation of Powers
“A Principle of Which We Were Ashamed”
The Three-Fifths Compromise
The Slave Trade
Fugitive Slaves
“The National Calamity”

An Overview of the Constitution


Preamble
Article 1: Congress

HAVE WE ACHIEVED THE


WHAT DO YOU THINK?
CONSTITUTION’S GOALS TODAY?
Article 2: The President
Article 3: The Courts
Article 4: Relations Between the States
Article 5: Amendments
Article 6: The Law of the Land
Article 7: Ratification
The Missing Articles

Ratification
The Anti-Federalists
The Federalists
Two Strong Arguments
A Very Close Vote

Changing the Constitution


The Bill of Rights
The Seventeen Amendments
The Constitution Today
INFO DATA Amend the Constitution Today? On
What Issue?
HOW STRICTLY SHOULD WE
WHAT DO YOU THINK?
INTERPRET THE CONSTITUTION?
Conclusion: Does the Constitution Still Work?
Chapter Summary
Key Terms
Study Questions

3 Federalism and Nationalism


BY THE NUMBERS Federalism

Forging Federalism
Who Holds Government Authority?
Advantages of State-Level Policy
The Advantages of National Policy
INFO DATA Regulatory Policies: Differ By State
PRESERVING LOCAL VALUES
WHAT DO YOU THINK?
OR PROMOTING CONSISTENT NATIONAL
POLICY?
How Federalism Works
The Constitution Sets the Ground Rules
The Constitution Empowers National Authority
The Constitution Protects State Authority
The Constitution Authorizes Shared Power
Dual Federalism (1789–1933)
Cooperative Federalism (1933–1981)
New Federalism
Progressive Federalism
Education
Healthcare
Federalism Today

Issues in Federalism
Unfunded Mandates
Drowned in the Bathtub? Reducing the Federal Government
Federalism in the Courts

Nationalism, American Style


The Rise of American Nationalism
America’s Weak National Government
Size
Authority
Independence

Conclusion: Who Are We?


Chapter Summary
Key Terms
Study Questions

4 Civil Liberties
BY THE NUMBERS Civil Liberties

The Rise of Civil Liberties


Civil Rights and Civil Liberties
The Purpose of Civil Liberties
The Slow Rise of Civil Liberties

Privacy
Penumbras and Emanations
Roe v. Wade

WHAT DO YOU THINK? IS THERE A RIGHT TO


PRIVACY?
Planned Parenthood v. Casey
Sex Between Consenting Adults
Clashing Principles

Freedom of Religion
The Establishment Clause
Free Exercise of Religion

MAY THE CHRISTIAN YOUTH


WHAT DO YOU THINK?
CLUB MEET IN SCHOOL?
Freedom of Speech
A Preferred Position
Political Speech
Symbolic Speech
Limits to Free Speech: Fighting Words
Limited Protections: Student Speech

WHAT DO YOU THINK? FREE SPEECH ON CAMPUS


Freedom of the Press
Prior Restraint
Obscenity
Libel

The Right to Bear Arms


A Relic of the Revolution?
The Palladium of All Liberties?

INFO DATA Guns On Campus: Concealed Carry?


The Rights of the Accused
The Fourth Amendment: Search and Seizure
The Fifth Amendment: Rights at Trials
The Sixth Amendment: The Right to Counsel
The Eighth Amendment: The Death Penalty

Terrorism, Non-Citizens, and Civil Liberties


Contacts with Forbidden Groups
Surveillance
The Rights of Non-Citizens

Conclusion: The Dilemma of Civil Liberties


Chapter Summary
Key Terms
Study Questions

5 The Struggle for Civil Rights

Winning Rights: The Political Process


Seven Steps to Political Equality
BY THE NUMBERS Civil Rights

How the Courts Review Cases


Suspect Categories
Quasi-Suspect Categories
Nonsuspect Categories

Race and Civil Rights: Revolt Against Slavery


The Clash over Slavery
Abolition
Economics
Politics
Dred Scott v. Sandford
The Second American Founding: A New Birth of Freedom?
Freedom Fails

The Fight for Racial Equality


Two Types of Discrimination
The Modern Civil Rights Campaign Begins
The Courts
The Civil Rights Movement
Congress and the Civil Rights Act
The Post Civil Rights Era
Affirmative Action in the Workplace

HIGHER EDUCATION AND


WHAT DO YOU THINK?
AFFIRMATIVE ACTION
Affirmative Action in Education

Women’s Rights
Suffrage
The Civil Rights Act of 1964
Equal Rights Amendment
The Courts
Progress for Women—But How Much?

Hispanics
Challenging Discrimination
The Politics of Immigration
Ancient Fears
Three Categories
Undocumented Individuals
Language Controversy: Speak English!
Political Mobilization

Asian Americans
Native Americans
The Lost Way of Life
Indians and the Federal Government
Social Problems and Politics

Groups Without Special Protection


People with Disabilities
Sexual Orientation and Gender Identity
INFO DATA Protection for Sexual Orientation and
Gender Identity: How Do Employment Laws
Vary by State?
The Fight for Civil Rights Goes On
Voting Rights Today
Economic and Social Rights Today
Health
Income
Incarceration

Conclusion: Civil Rights . . . By the People


Chapter Summary
Key Terms
Study Questions

PART II POLITICAL BEHAVIOR

6 Public Opinion and Political Participation


BY THE NUMBERS Public Opinion and Political Participation

Sources of Public Opinion


Political Socialization
Parents and Friends
Education
Gender
Race
Religion
Life Events
Party
Self-Interest: Voting Our Pocketbooks
Elite Influence
Wars and Other Focusing Events

Measuring Public Opinion


Polling Bloopers
Polling 101
The Random Sample
Sampling frame
Refining the Sample
Timing

Talking Politics Types of Polls


Wording
Lies, Damn Lies, and Polls
Technology and Error
Sampling Error and Response Bias
Do Opinion Surveys Influence Us?

Public Opinion in a Democracy


Ignorant Masses
The Rational Public
Governing By the People

HOW CLOSELY SHOULD


WHAT DO YOU THINK?
ELECTED OFFICIALS FOLLOW PUBLIC
OPINION?
Do the People Know What They Want?
How Do the People Communicate Their Desires?
Do Leaders Respond to Public Opinion?

Getting Involved: Political Participation


Traditional Participation
Voting
Electoral Activities
Voice
Civic Voluntarism
Direct Action

WHAT DO YOU THINK? WOULD YOU HAVE


PROTESTED?
The Participation Puzzle

Why Do People Get Involved?


Background: Age, Wealth, Education, and Race
Age
Wealth
Education
Race
Friends and Family
Community
Political Mobilization

INFO DATAPublic Rallies: What Issues Draw


People to Protest?
Government Beneficiaries
Historical Context

Talking Politics How Mobilizers See Us


What Discourages Political Participation?
Alienation
Institutional Barriers
Complacency
Shifting Mobilization Patterns

New Avenues for Participation: The Internet, Social


Media, and Millennial Participation
Scenario 1: Rebooting Democracy
Scenario 2: More Hype and Danger than Democratic Renaissance
Does Social Media Increase Political Participation?
How the Millennial Generation Participates

Conclusion: Government by the People


Chapter Summary
Key Terms
Study Questions

7 Media, Technology, and Government


BY THE NUMBERS The Media

Media and American Democracy


Providing Information
Watching Officials
Shaping the Policy Agenda

U.S. Media Today: Traditional Formats Are Declining


Newspapers and Magazines: Rise and Decline
Radio Holds Steady
Television: From News to Infotainment
The Rise of Cable
Infotainment

New Media Rising


Is the Media Biased?
Are Reporters Politically Biased?
Profits Drive the News Industry
Drama Delivers Audiences
Investigative “Bias”

WHAT DO YOU THINK? IS THE MEDIA OBJECTIVE?


The Fairness Bias

How Governments Shape the Media


The First Amendment Protects Print Media from Regulation
Regulating Broadcasters
Protecting Competition
INFO DATA Media Consolidation: Who Produces,
Distributes, and Owns the Media?
Media Around the World
Government-Owned Stations
Censorship
American Media in the World

Media in Context: War, Terrorism, and U.S. Elections


Covering Wars and Terrorism
The Campaign as Drama
Candidate Profiles
Conclusion: At the Crossroads of the Media World
Chapter Summary
Key Terms
Study Questions

8 Campaigns and Elections

How Democratic Are U.S. Elections?


BY THE NUMBERS Campaigns and Elections

Frequent and Fixed Elections


Over 520,000 Elected Officials
Barriers to Voting

ELECTED VERSUS
WHAT DO YOU THINK?
APPOINTED POSITONS?
Financing Campaigns: The New Inequality?
Too Much Money?
Democracy for the Rich?
Major Donors: Easier to Give

INFO DATA Money in Elections: The New Rules


Presidential Campaigns and Elections
Who Runs for President?
The Three Phases of Presidential Elections
Winning the Nomination

WHAT DO YOU THINK? WHY IOWA AND NEW


HAMPSHIRE?
Organizing the Convention
The General Election
Winning Presidential Elections
Economic Outlook
Demographics
War and Foreign Policy
Domestic Issues
The Campaign Organization
Parties Matter
The Electoral College and Swing States
That Elusive Winning Recipe
Predicting Presidential Elections

Congressional Elections
Candidates: Who Runs for Congress?
The Power of Incumbency
Patterns in Congressional Elections
Redrawing the Lines: The Art of Gerrymandering
Nonpartisan Districting and Minority Representation

How to Run for Congress


Key 1: Money
Key 2: Organization
Key 3: Strategy

Talking Politics Campaign Lingo


Key 4: Message

Conclusion: Reforming American Elections


Chapter Summary
Key Terms
Study Questions

9 Interest Groups and Political Parties

The Many Roles Interest Groups Play


BY THE NUMBERS

Informing Members
Communicating Members’ Views
Mobilizing the Public
What Interest Groups Do for Democracy
Types of Interest Groups
Economic Groups
Citizen or Public Interest Groups
Intergovernmental and Reverse Lobbying

Interest Groups and the Federal Government


The Multiple Roles of Lobbyists
Researchers
Witnesses
Position Takers
Coalition Builders
Social Butterflies
Iron Triangles
Rise of the Issue Network

Talking Politics Washington Lobbying


Interest Groups and the Courts
Lobbying on Judicial Confirmations
Filing Amicus Curiae (literally, Friend of Court) Briefs
Sponsoring Litigation
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Mouth-Frame.—In its broad outlines there is practically no variation
in this organ throughout the group, but in respect of the spines,
which are borne on the flanks of the jaws (mouth-papillae) and on
their apices (teeth and tooth-papillae) there is very great variation.
Teeth are always present. Mouth-papillae are very frequently
present, tooth-papillae are rarer, and it is only in a restricted number
of genera (Ophiocoma and its allies) that both mouth-papillae and
tooth-papillae are present at the same time.

Fig. 216.—A portion of an arm of Ophiohelus umbella, near the distal extremity,
treated with potash to show the skeleton, × 55. The vertebrae are seen to
consist of two curved rods united at their ends. The triangular side-plates
bear a row of movable hooks which articulate with basal outgrowths of the
plate. (After Lyman.)

Skeleton of the Disc.—This is typically composed of a mosaic of


plates of different sizes, but in some cases (Ophiomyxa, most
Streptophiurae, and Cladophiurae) these, with the exception of the
radials and genitals, are entirely absent, and the disc is then quite
soft and covered with a columnar epithelium, the persistent
ectoderm. Even the scuta buccalia may disappear. Radial shields
are absent in Ophiohelus. In many cases (Ophiothrix and
Ophiocoma) all the dorsal plates except the radials are concealed
from view by a covering of small spines. In some genera
(Ophiopyrgus) there are five large plates in the centre of the upper
part of the disc, which have been termed "calycinals" from a
mistaken comparison with the plates forming the cup or calyx of the
Pelmatozoa, but there is no connexion between the two sets of
structures.
The madreporite is usually quite rudimentary, but in Cladophiurae
there may be five madreporites, each with about 200 pores, and, of
course, five stone-canals.

The number of genital organs varies very much. In the small


Amphiura squamata there are two gonads, an ovary and a testis,
attached to each bursa, but in the larger species there may be very
many more.

We follow Bell's classification,[463] according to which the


Ophiuroidea are divided, according to the manner in which the
vertebrae move on one another (cf. Fig. 210), into three main orders,
since these movements are of prime importance in their lives.

(1) Streptophiurae, in which the faces of the vertebrae have


rudimentary knobs and corresponding depressions, so that the arms
can be coiled in the vertical plane. These are regarded as the most
primitive of Ophiuroidea.

(2) Zygophiurae, in which the vertebral faces have knobs and pits
which prevent their coiling in a vertical plane.

(3) Cladophiurae, in which the arms can be coiled as in (1) and are
in most cases forked. No teeth; the arm-spines are papillae, the
covering plates of the arms are reduced to granules.

Order I. Streptophiurae.
This is not a very well defined order; it includes a few genera
intermediate in character between the Cladophiurae and the
Zygophiurae, and believed to be the most primitive Ophiuroids living.
It is not divided into families. The vertebrae have rudimentary
articulating surfaces, there being two low bosses and corresponding
hollows on each side, and so they are capable of being moved in a
vertical plane, as in the Cladophiurae; the arms never branch, and
further, they always bear arm-spines and lateral arm-plates at least.
No species of this order are found on the British coast, but
Ophiomyxa pentagona, in which the dorsal part of the disc is
represented only by soft skin, is common in the Mediterranean.

Ophioteresis is devoid of ventral plates on the arms, and appears to


possess an open ambulacral groove, though this point has not been
tested in sections. Ophiohelus and Ophiogeron have vertebrae in
which traces of the double origin persist (see p. 491).

Order II. Zygophiurae.


This group includes all the common and better-known British forms.
They are divided into five families, all of which are represented in
British waters.

Fig. 217.—Aboral view of Ophioglypha (Ophiura) bullata. × 3. (From Wyville


Thomson.)
Fam. 1. Ophiolepididae.[464]—Arm inserted in a definite cleft in the
disc, or (expressing the same fact in another way) the interradial
lobes out of which the disc is composed are not completely united.
Radial shields and dorsal plates naked. Arm-spines smooth and
inserted on the posterior border of the lateral arm-plates.

Fig. 218.—Oral view of Ophioglypha (Ophiura) bullata. × 5. (From Wyville


Thomson.)

This family includes all the Brittle Stars of smooth porcelanous


aspect and provided with only short spines. Forbes[465] called them
Sand-stars, since their short spines render these animals incapable
of burrowing or of climbing well, and hence they appear to move
comparatively rapidly over firm ground, sand, gravel, or muddy sand,
and they are active enough to be able to capture small worms and
Crustacea. The prey is seized by coiling one of the arms around it.

One genus, Ophiura, is fairly common round the British coast, being
represented by O. ciliaris and O. albida; the former is the commoner.
An allied species dredged by H.M.S. "Challenger" is represented in
Figs. 217 and 218.

Ophiomusium (Fig. 219) is a very peculiar genus. The mouth-


papillae on each side of each mouth-angle are confluent, forming a
razor-like projection on each side of each mouth-angle (Fig. 220).
The arms are short, and the podia are only developed at the bases
of the arms. Ophiopyrgus has the dorsal surface raised into a conical
elevation protected by a central plate surrounded by five large
plates.

Fig. 219.—Aboral view of Ophiomusium pulchellum. × 7. (From Wyville


Thomson.)

In the remaining four families the arms are inserted on the under
surface of the disc; in other words, the interradial lobes which make
up the disc have completely coalesced dorsally; and the spines
stand out at right angles to the arm.

Fam. 2. Amphiuridae.—Mouth-papillae present, but no tooth-


papillae; radial shields naked; small scuta buccalia.

The most interesting Brittle Star belonging to this family is Amphiura


squamata (elegans), a small form, with a disc about ¼ inch in
diameter covered with naked plates. It is hermaphrodite and
viviparous, the young completing their development inside the
bursae of the mother. Occasionally the whole disc, with the
exception of the mouth-frame, is thrown off and regenerated. This
appears to be a device to enable the young to escape. Three other
species of Amphiura are found in British waters.

Fig. 220.—Oral view of Ophiomusium pulchellum. × 7. (From Wyville Thomson.)

Ophiactis is another genus belonging to this family, distinguished


from Amphiura by its shorter arms and smoother arm-spines. It lives
in the interstices of hard gravel. The British species, O. balli,
presents no special features of interest, but the Neapolitan O. virens
is an extraordinary form. It has six arms, three of which are usually
larger than the other three, for it is always undergoing a process of
transverse division, each half regenerating the missing part. It has
from 1 to 5 stone-canals, the number increasing with age; numerous
long-stalked Polian-vesicles in each interradius, and in addition a
number of long tubular canals which spring from the ring-canal, and
entwine themselves amongst the viscera.[466] All the canals of the
water-vascular system, except the stone-canals, contain non-
nucleated corpuscles, carrying haemoglobin,[467] the respiratory
value of which compensates for the loss of the genital bursae, which
have entirely disappeared.

Ophiopholis is distinguished from the foregoing genera by the


granular covering of its dorsal plates; whilst in Ophiacantha these
granules develop into prominent spinelets, and the arm-spines are
also thorny. Ophiopholis aculeata occurs in swarms in the branches
of the Firth of Clyde, and presents a most remarkable series of
variations in colour. Ophiopsila is a closely allied form, distinguished
by its large peristomial plates.

Fig. 221.—Oral view of Ophiacantha chelys. × 4. (From Wyville Thomson.)

Fam. 3. Ophiocomidae.—Both mouth-papillae and tooth-papillae


are present;[468] the arm-spines are smooth, and the disc is covered
with granules.

Ophiocoma nigra is the only common British representative of this


family. In this species the plates of the dorsal surface are completely
hidden from view by a covering of granules. Ophiarachna.

Fam. 4. Ophiothricidae.—Tooth-papillae alone present, mouth-


papillae absent; arm-spines roughened or thorny.

This family is represented only by Ophiothrix fragilis, which is


perhaps the most abundant of all British Ophiuroids, and has been
selected as the type for special description.

The back is covered with spinules, having, however, the triangular


radial plates bare. This produces a contrast-effect, which suggested
the name pentaphyllum, formerly used by some naturalists for the
species. It occurs in swarms, and presents variations in colour nearly
as marked as those of Ophiopholis. Ophiopteron is probably a
swimming Ophiuroid, as the lateral spines of each segment of the
arm are connected by a web of skin.

Order III. Cladophiurae.

Fig. 222.—Aboral view of young Astrophyton linckii, slightly enlarged. (From


Wyville Thomson.)

These, like the Streptophiurae, have the power of rolling the arms in
a vertical plane, but the articulating surfaces of the vertebrae are
well-developed and saddle-shaped. The dorsal surface of the disc
and arms is covered with a thick skin with minute calcifications.
Upper-arm plates wanting. Radial plates always present, though
occasionally represented by lines of scales. The order is divided into
three families, two of which are represented in British waters.

Fam. 1. Astroschemidae.—Arms unbranched. Astronyx is


comparatively common in the sea-lochs of Scotland. There are a
series of pad-like ridges on the arms, representing the side-plates
and bearing the spines. Astroschema.
Fam. 2. Trichasteridae.—Arms forked only at the distal ends.
Trichaster, Astrocnida.

Fam. 3. Euryalidae.—Arms forked to their bases. Gorgonocephalus


is occasionally taken in deep water off the north coast of Scotland. In
it the arms repeatedly fork, so that a regular crown of interlacing
arms is formed. The animal obviously clings to external objects with
these, for it is often taken in fishermen's nets with its arms coiled
around the meshes. The genital bursae are said to be represented
by slits which open directly into the coelom. (Lyman describes the
coelom as divided into ten compartments by radiating septa; it is
possible—even probable—that these are really the bursae.) An allied
species is common in the Bay of Fundy, being found in
comparatively shallow water. Astrophyton (Fig. 222) is closely allied
to Gorgonocephalus, differing only in trifling points. It is doubtful
whether the separation of these two genera is justified.

Fossil Ophiuroidea.—The Ophiuroidea are rather sparsely


represented among fossils, but in the Silurian and Devonian a series
of very interesting forms occur which are intermediate in character
between Starfish and Brittle Stars, and which were therefore in all
probability closely allied to the common ancestors of modern
Ophiuroids and Asteroids. Jaekel[469] has recently added largely to
our knowledge of these primitive forms, and has described a number
of new genera. Thus Eophiura from the Lower Silurian has an open
ambulacral groove, and the vertebrae are represented by an
alternating series of quadrate ossicles, each deeply grooved on its
under surface for the reception of the tentacle, which was not yet (as
in modern forms) enclosed in the vertebra. The lateral or
adambulacral plates extended horizontally outwards, and each bore
a series of spines at its outer edge.

A remarkable fact is that where the halves of the vertebrae (i.e. the
ambulacral ossicles) diverge in order to form the mouth-angles, no
less than five or six vertebrae are thus affected, instead of only two
as in modern forms. The actual "jaw," however, seems, as in modern
forms, to consist only of the first adambulacral fused to the second
ambulacral, so that instead of concluding with Jaekel that the "jaws"
of modern forms result from the fusion of five or six vertebrae, a
conclusion which would require that a number of tentacles had
disappeared, we may suppose that the gaping "angles" of these old
forms have, so to speak, healed up, except at their innermost
portions.

In Bohemura, which belongs to a somewhat younger stratum, the


structure is much the same, but the groove in the ambulacral ossicle
for the tentacle has become converted into a canal, and the
ambulacral groove itself has begun to be closed at the tip of the arm
by the meeting of the adambulacrals.

In Sympterura, a Devonian form described by Bather,[470] the two


ambulacral plates of each pair have thoroughly coalesced to form a
vertebra, but there is still an open ventral groove, and no ventral
plates.

In the Trias occurs the remarkable form Aspidura, which had short
triangular arms, in which the tentacle pores were enormous and the
ventral plates very small. The radial plates formed a continuous ring
round the edge of the disc. Geocoma from the Jurassic is a still more
typical Ophiuroid; it has long whip-like arms, and the dorsal skeleton
of the disc is made of fifteen plates, ten radials, and five interradials.
In the Jurassic the living genus Ophioglypha, appears.

The Cladophiurae are represented already in the Upper Silurian by


Eucladia, in which, however, the arms branch not dichotomously, as
they do in modern forms, but monopodially. There is a large single
madreporite.

Onychaster, with unbranched arms, which occurs in the


Carboniferous, is a representative of the Streptophiurae.
It will therefore be seen that the evolution of Ophiuroidea must have
begun in the Lower Silurian epoch. The Streptophiurae are a few
slightly modified survivors of the first Ophiuroids. By the time the
Devonian period had commenced, the division of the group into
Zygophiurae and Cladophiurae had been accomplished.

CHAPTER XVIII

ECHINODERMATA (CONTINUED): ECHINOIDEA = SEA-URCHINS

CLASS III. ECHINOIDEA


The Sea-urchins or Echinoidea (Gr. ἐχῖνος, Hedgehog or Sea-
urchin), which constitute the third class of the Eleutherozoa, have
derived both their popular and scientific names from the covering of
long spines with which they are provided. At first sight but little
resemblance is to be discerned between them and the Starfish and
Brittle Stars. They are devoid of any outgrowths that could be called
arms; their outline is generally either circular or that of an equilateral
pentagon, but as their height is almost always smaller than their
diameter, they are never quite spherical; sometimes it is so small
that the animals have the form of flattened discs.

All doubt as to the relationship of the Echinoidea to the Starfish is at


once dispelled in the mind of any one who sees one of the common
species alive. The surface is beset with delicate translucent tube-
feet, terminated by suckers resembling those of Starfish, although
capable of much more extension. The animal throws out these
organs, which attach themselves by their suckers to the substratum
and so pull the body along, whilst the spines are used to steady it
and prevent it from overturning under the unbalanced pull of the
tube-feet. When moving quickly the animal walks on its spines, the
tube-feet being little used. The tube-feet are distributed over five
bands, which run like meridians from one pole of the animal to the
other. These bands are termed "radii," and they extend from the
mouth, which is situated in the centre of the lower surface, up to the
neighbourhood of the aboral pole. The radii must be compared to the
ambulacral grooves on the oral surface of the arms of Starfish, and
hence in Urchins the aboral surfaces of the arms have, so to speak,
been absorbed into the disc, so that the oral surfaces have become
bent in the form of a semicircle. The radii are separated from one
another by meridional bands called "interradii," which correspond to
the interradial angles of the disc of a Starfish and to the sides of its
arms. The small area enclosed between the upper terminations of
the radii is called the "periproct," and this corresponds to the entire
dorsal surface of the Starfish, including that of the arms.

One of the commonest species of British Sea-urchin is Echinus


esculentus. In sheltered inlets, such as the Clyde, it is often left
exposed by the receding tide, whilst everywhere on the coast in
suitable localities it may be obtained by dredging at moderate depths
on suitable ground. In the Clyde it is easy to observe the habits of
the animal through the clear still water. It is then seen to frequent
chiefly rocky ground, and to exhibit a liking for hiding itself in
crevices. Often specimens will be seen clinging to the rock by some
of their tube-feet, and, as it were, pawing the under surface of the
water with the others. In the Clyde it feeds chiefly on the brown
fronds of Laminaria, with which the rocks are covered. In more
exposed situations, such as Plymouth Sound, it does not occur in
shallower water than 18 to 20 fathoms. At this depth it occurs on a
rocky ridge; but in 1899, after a south-west gale, all the specimens
had disappeared from this ridge, showing at what a depth wave
disturbance is felt.

A full-grown specimen is as large as a very large orange; its under


surface is flattened, and it tapers somewhat towards the aboral pole.
The outline is that of a pentagon with rounded angles. The spines in
Echinus esculentus are short in comparison to the diameter of the
body, and this is one of the characteristics of the species.
The animal is provided with a well-developed skeleton, consisting of
a mail of plates fitting closely edge to edge, and carrying the spines.
This cuirass bears the name "corona" (Fig. 227). It has two
openings, an upper and a lower, which are both covered with flexible
skin. The upper area is known as the "periproct" (Fig. 227, 2); it has
in it small isolated plates, and the anus, situated at the end of a small
papilla, projects from it on one side of the centre. The lower area of
flexible skin surrounds the mouth, and is called the "peristome" (Fig.
229), though it corresponds to considerably more than the peristome
of Asteroidea. In the mouth the tips of the five white chisel-like teeth
can be seen.

The plates forming the corona are, like all the elements of the
skeleton of Echinodermata, products of the connective tissue which
underlies the ectoderm, which in Echinoidea remains in a fully
developed condition covering the plates, and does not, as in
Ophiuroidea, dry up so as to form a mere cuticle. The ectoderm
consists of the same elements as that of Asteroidea, viz. delicate
tapering sense-cells with short sense-hairs, somewhat stouter
supporting cells and glandular cells. It is everywhere underlaid by a
plexus of nerve fibrils, which, in part, are to be regarded as the basal
outgrowths of the sense-cells and partly as the outgrowths of a
number of small bipolar ganglion-cells, found intermixed with the
fibres.

Fig. 223.—Aboral view of Echinus esculentus. × ½. (After Mortensen.)


Just as the muscular arm has been the determining factor in the
structure of the Ophiuroidea, so the movable spine has been the
leading factor in the evolution of Echinoidea. The spines have cup-
shaped basal ends, which are inserted on special projections of the
plates of the skeleton called tubercles. The tubercle is much larger
than the cup, and hence the spine has a great range of possible
motion. The spines differ from those of Starfish and Brittle Stars in
being connected with their tubercles by means of cylindrical sheaths
of muscle fibres, by the contraction of which they can be moved in
any direction. The muscles composing the sheath consist of an outer
translucent and an inner white layer. The former are easily
stimulated and soon relax; they cause the movements of the spines.
The latter require stronger stimulation, but when aroused respond
with a prolonged tetanus-like contraction, which causes the spines to
stand up stiffly in one position; these muscles can be torn across
sooner than forced to relax. Uexküll[471] has appropriately named
them "block musculature." These sheaths, like everything else, are
covered with ectoderm, which is, however, specially nervous, so that
we may say that the muscular ring is covered by a nerve-ring from
which stimuli are given off to the muscles.

The spines are, speaking generally, of two sizes, the larger being
known as "primary spines" and the smaller as "secondary." In many
Echinoidea these two varieties are very sharply contrasted, but in
Echinus esculentus there is not such a great difference in length,
and intermediate kinds occur. The forest of spines has an
undergrowth of pedicellariae. All Echinoidea possess pedicellariae,
which are much more highly developed than those of any Asteroid.
With few exceptions all the pedicellariae of Echinoidea possess
three jaws and a basal piece. This latter is, however, drawn out so as
to form a slender rod, which articulates with a minute boss on a plate
of the skeleton.

Of these pedicellariae there are in E. esculentus four varieties, viz.


(1) "tridactyle" (Fig. 225, C; Fig. 226, B): large conspicuous
pedicellariae with three pointed jaws, each armed with two rows of
teeth on the edges. There is a flexible stalk, the basal rod reaching
only half way up. These are scattered over the whole surface of the
animal.

(2) "Gemmiform" (Fig. 225, A, B; Fig. 226, A), so called from the
translucent, almost globular head. The appearance of the head is
due to the fact that there is on the outer surface of each jaw a sac-
like gland developed as a pouch of the ectoderm. From it are given
off two ducts which cross to the inner side of the blades and, uniting
into one, run in a groove to near the tip. The gland secretes a
poisonous fluid. The basal rod reaches up to the jaws, so that this
form of pedicellaria has a stiff stalk. On the inner side of each blade,
near the base, there is a slight elevation (Fig. 225, B, s), consisting
of cells bearing long cilia; this is a sense-organ for perceiving
mechanical stimuli. The gemmiform pedicellariae are particularly
abundant on the upper surface of the animal.

Fig. 224.—View of the apical region of Echinus esculentus, showing spines and
pedicellariae; drawn from the living specimen, × 3. a, Anus; g.p, genital
pore; i, interradius; mp, madreporite; per, periproct; p.gemm, gemmiform
pedicellaria; pod, podia; p.trid, tridactyle pedicellaria; p.trif, trifoliate
pedicellaria; r, radius; t.t, pore for terminal tentacle of the radial water-
vascular canal.
(3) "Trifoliate" (Fig. 225, E; Fig. 226, D): these are very small
pedicellariae, in which the jaws are shaped like leaves with the broad
end projecting outwards. They are scattered over the whole surface
of the body.

(4) "Ophicephalous" (Fig. 225, D; Fig. 226, C): pedicellariae in which


the jaws have broad rounded distal ends fringed with teeth; these
ends bear a resemblance to a snake's head, whence the name. The
bases are also broad and thin, with a strong median rib and a
peculiar semicircular hoop beneath the spot where they articulate
with one another. The three hoops of the three jaws work inside each
other in such a way as to cause the jaws to have a strong grip and to
be very difficult to dislocate from their mutual articulation.

The ophicephalous pedicellariae are in Echinus the most abundant


of all; and they alone extend on to the peristome, where a special
small variety of them is found.

A thorough investigation of the functions and reactions of the


pedicellariae has quite recently been made by von Uexküll.[472] He
showed, first of all, that there is a nervous centre in the stalk of each
pedicellaria (see below), which causes the organ to incline towards a
weak stimulus, but to bend away from a stronger stimulus. In the
head there is an independent nervous centre, which regulates the
opening and closing of the valves, and causes these to open on
slight stimulus and close when a stronger one is applied. The
amount of stimulus necessary to cause the pedicellariae to retreat
varies with the kind of pedicellariae, being least with the tridactyle
and most with the gemmiform, so that when a chemical stimulus,
such as a drop of dilute ammonia, is applied to the skin, the
tridactyle pedicellariae may be seen to flee from and the gemmiform
to approach the point of stimulation. In a living Sea-urchin, if the
attempt is made to seize the tridactyle pedicellariae they will evade
the forceps, but the ophiocephalous are easy to catch.
The tridactyle pedicellariae open with the very slightest mechanical
stimulus and close with rather greater mechanical stimuli or with
exceedingly slight chemical ones. Uexküll calls them "Snap-
pedicellariae," and their function is to seize and destroy the minute
swimming larvae of various sessile parasitic animals, which would
otherwise settle on the delicate exposed ectoderm of the Sea-urchin.

The gemmiform pedicellariae are brought into action when a more


serious danger threatens the Sea-urchin, such as an attack of a
Starfish. The corrosive chemical influence, which it can be proved
exudes not only from the stomach but even from the tube-feet of the
Starfish, causes the gemmiform pedicellariae to approach and open
widely. When the foe approaches so closely as to touch the sense-
organs (Fig. 225, B, s) situated on the inner side of the valves of
these pedicellariae, the blades close violently, wounding the
aggressor and causing its juice to exude, thus producing a renewed
and severe chemical stimulation which irritates the poison glands
and causes the poison to exude. The virulence of the poison may be
gauged from the fact that the bite of a single gemmiform pedicellaria
caused a frog's heart to stop beating.

Fig. 225.—The pedicellariae of Echinus acutus, drawn from a living specimen. A,


gemmiform pedicellaria, closed. B, gemmiform pedicellaria, open; g, poison
gland; s, sense-organ, × 3. C, tridactyle pedicellaria, × 6. D, ophicephalous
pedicellaria, × 9. E, trifoliate pedicellaria, × 12; a (in all figures), axial rod of
the stalk. (After Uexküll.)

Prouho[473] has described a combat between a Sea-urchin and a


Starfish. When the latter approached, the spines of the Sea-urchin
diverged widely (strong form of reaction to chemical stimulus),
exposing the gemmiform pedicellariae. These at once seized the
tube-feet of the enemy and the Starfish retreated, wrenching off the
heads of these pedicellariae; then the Starfish returned to the attack
and the same result followed, and this was repeated till all the
pedicellariae were wrenched off, when the Starfish enwrapped its
helpless victim with its stomach.

The minute trifoliate pedicellariae are brought into play by any


prolonged general irritation of the skin, such as bright light or a rain
of particles of grit or mud. They have the peculiarity that not all the
blades close at once, so that an object may be held by two blades
and smashed by the third. They may be seen in action if a shower of
powdered chalk is poured on the animal, when they seize the
particles and by breaking up any incipient lumps reduce the whole to
an impalpable powder, which the cilia covering the skin speedily
remove. In thus assisting in the removal of mechanical "dirt" they
earn the name which Uexküll has bestowed on them, of "cleaning
pedicellariae."

Fig. 226.—Views of a single blade of each kind of pedicellaria. A, blade of


gemmiform pedicellaria of Echinus elegans; g, groove for duct of poison
gland; B, blade of tridactyle pedicellaria of the same species; C, blade of
ophicephalous pedicellaria of the same species; r, ring for clamping this
blade to the other blades; D, blade of trifoliate pedicellaria of E. alexandri.
(After Mortensen.)

The ophicephalous pedicellariae, with their powerful bull-dog grip,


assist in holding small animals, such as Crustacea, till the tube-feet
can reach them and convey them to the mouth.

The number and variety of the pedicellariae, then, is an eloquent


testimony to the dangers to which the soft sensitive skins of the Sea-
urchin and other Echinodermata are exposed, and afford
confirmatory evidence in support of the view expressed above, that
the method adopted to defend the skin was one of the great
determining features which led to the division of the Asteroidea into
different races.

Fig. 227.—Dried shell of Echinus esculentus, showing the arrangement of the


plates of the corona. × 1. 1, The anus; 2, periproct, with irregular plates; 3,
the madreporite; 4, one of the other genital plates; 5, an ocular plate; 6, an
interambulacral plate; 7, an ambulacral plate; 8, pores for protrusion of the
tube-feet; 9, tubercles of the primary spines, i.e. primary tubercles.

The corona consists of five radial or "ambulacral" bands of plates


and five interradial, or as they are usually termed, "interambulacral"
bands of plates—ten in all. Each of the ten consists of two vertical
rows of plates throughout most of its extent, and each plate is
studded with large bosses, or "primary tubercles" for the primary
spines, smaller bosses called "secondary tubercles" for the
secondary spines, and finally, minute elevations called "miliary
tubercles" for the pedicellariae.
Fig. 228.—The so-called calyx and the periproct of Echinus esculentus. × 4. 1,
Genital plates with genital pores; 2, ocular plates with pores for terminal
tentacles of the radial water-vascular canals; 3, madreporite; 4, periproct
with irregular plates; 5, anus. (After Chadwick.)

Even in the dried skeleton, however, the ambulacral plates can be


discriminated from the interambulacral by the presence of pores to
permit the passage of the tube-feet. These pores are arranged in
pairs, and each pair corresponds to a single tube-foot, since the
canal connecting the ampulla with the external portion of the tube-
foot is double in the Echinoidea. In Echinus esculentus there are
three pairs of such pores in each plate, in Strongylocentrotus
droëbachiensis four pairs. The ambulacral plate is really made up of
a series of "pore-plates," each carrying a single pair of pores, and
these become united in threes in Echinus and fours in
Strongylocentrotus, while in primitive forms like the Cidaridae they
remain separate. Each ambulacral and interambulacral area ends at
the edge of the periproct with a single plate. The plate terminating
the ambulacral band is pierced by a single pore for the exit of the
median tentacle, which, as in Asteroids, terminates the radial water-
vascular canal. Thus the aboral end of the radius in an Echinoid
corresponds to the tip of the arm in an Asteroid. The plate is termed
"ocular," because the terminal tentacle has a mass of pigmented
cells at its base; but no eye-cups can be seen, and there is no
evidence that this spot is specially sensitive to light. Species which
show special sensitiveness to light have often a large number of
what we may perhaps term secondary eyes. The plate terminating
the interambulacral series is termed the "genital plate," because it is
pierced by the duct of one of the five genital organs. One of the

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