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Chapter 8 Mechanisms and Inhibitors

Matching Questions
Use the following to answer questions 1–10.

Choose the correct answer from the list below. Not all of the answers will be used.
a) hydrolysis
b) affinity label
c) tyrosinase
d) chymotrypsin
e) pepsin
f) noncompetitive
g) uncompetitive
h) heterotropic
i) approximation and orientation
j) acid–base
h) competitive

1. ____________: An enzyme that temporarily undergoes covalent catalysis as part of its

mechanism.
Ans: d
Section: 8.3

2. ____________: The type of reaction catalyzed by proteases.

Ans: a
Section: 8.3

3. ____________: A protease enzyme with a low pH optimum.

Ans: e
Section: 8.2

4. ____________: The type of catalysis in which two substrates are brought into close proximity.
Ans: j
Section: 8.1

5. ____________: A molecule that is also known as a substrate analog.

Ans: b
Section: 8.2

6. ____________: The inhibitor which binds only to the ES complex and lowers the Vmax and KM.
Ans: g
Section: 8.2

7. ____________: The enzyme inhibition that can be overcome by increasing the concentration of
substrate.
Ans: k
Section: 8.2
Chapter 8 Mechanisms and Inhibitors 2

8. ____________: A type of catalysis where the proton donor is not water.

Ans: i
Section: 8.1

9. ____________: A type of enzyme inhibitor where KM is unaltered.

Ans: h
Section: 8.2

10. ____________: An enzyme that is part of a pigment formation pathway and has a low optimum
temperature.
Ans: c
Section: 8.2

Fill-in-the-Blank Questions
11. An enzyme catalyst mechanism that uses a metal cation to stabilize a negative charge in the
active site is ________________.
Ans: metal ion catalysis Section: 8.1

12. A ________________ catalytic mechanism that forces two substrates into an appropriate three-
dimensional arrangement for the reaction to occur.
Ans: catalysis by approximation and orientation Section: 8.1

13. The antibiotic penicillin is an example of a(n) ________________ inhibitor.

Ans: irreversible Section: 8.2

14. In conducting an experiment with a new drug, you find that regardless of the concentration of
substrate, the drug is able to inhibit the enzyme activity. You are likely to not have a(n)
________________ type of inhibitor.
Ans: competitive Section: 8.2

15. An uncompetitive inhibitor will have two ________________ lines on a double-reciprocal plot.
Ans: parallel Section: 8.2

16. A ________________ inhibitor binds irreversibly to the active site of an enzyme.

Ans: suicide Section: 8.2

17. The ________________ stabilizes the tetrahedral intermediate of the hydrolysis of a peptide
bond by chymotrypsin.
Ans: oxyanion hole Section: 8.3

18. A(n) ________________ inhibitor has a structure similar to the substrate and reversibly binds to
the active site of the enzyme.
Ans: competitive Section: 8.2

19. The straight-line kinetic plot of 1/V0 versus 1/S is called a ________________.
Ans: Lineweaver–Burk plot, or double-reciprocal plot Section: 8.2

20. The mechanism of chymotrypsin involves the formation of an unstable ________________-

shaped intermediate that is stabilized by the oxyanion hole.
Ans: tetrahedral Section: 8.3
Chapter 8 Mechanisms and Inhibitors 3

Multiple-Choice Questions

21. What conclusion can be drawn concerning an inhibitor if the KM is the same in the presence and
absence of the inhibitor?
A) The inhibitor binds to the substrate.
B) The inhibitor has a structure that is not very similar to the substrate.
C) The inhibitor forms a reversible covalent bond with the enzyme.
D) The inhibitor binds to the same active site as the substrate.
E) The Vmax is larger in the presence of the inhibitor.
Ans: B Section: 8.2

22. Which type(s) of inhibition can be reversed?

A) competitive
B) noncompetitive
C) uncompetitive
D) All of the above.
E) None of the above.
Ans: D Section: 8.2

23. In this type of inhibition, the inhibitor can only bind to the ES complex to form an ESI complex.
A) competitive
B) noncompetitive
C) irreversible
D) uncompetitive
E) None of the above.
Ans: D Section: 8.2

24. Which amino acids in chymotrypsin are found in the active site and are participants in substrate
cleavage?
A) his, ser, asp
B) his, ser
C) asp, lys
D) lys, arg
E) his, ser, arg
Ans: A Section 8.3

25. How is specificity determined by chymotrypsin?

A) by an interaction of the active site amino acids with the substrate
B) by a binding of the N-terminus amino acid at the active site
C) by a covalent binding of a his residue to the substrate
D) by a conformational change upon the binding of substrate
E) by a binding of the proper amino acid into a deep pocket on the enzyme
Ans: E Section: 8.3
Chapter 8 Mechanisms and Inhibitors 4

26. Where does cleavage of the scissile bond by chymotrypsin occur?

A) between a his and a ser amino acid
B) on the N-terminal side of a phe or trp residue
C) on the C-terminal side of a phe or trp residue
D) at the N-terminal amino acid
E) on the C-terminal side of an arg or lys amino acid
Ans: C Section: 8.3

27. A protein that is optimally active at neutral pH is likely to have which in the active site?
A) the side chains of aspartate and glutamine
B) two histidine amino acid side chains
C) a glycine amino acid
D) one or more carboxyl groups
E) two amino groups
Ans: A Section: 8.2

28. The metal most commonly found at the active site of metalloproteases is:
A) zinc.
B) calcium.
C) selenium.
D) magnesium.
E) sodium.
Ans: A Section: 8.1

29. Binding of a water molecule to the zinc ion induces:

A) a hydronium ion to form.
B) a large conformation change in the binding site.
C) ionization of a his residue, which functions as a strong nucleophile.
D) a lowered pKa for water, which leads to the formation of a zinc-bound hydroxide ion.
E) an altered KM value.
Ans: D Section: 8.1

Short-Answer Questions
30. How are the types of inhibition kinetically distinguishable?
Ans: Competitive inhibition can be overcome by the presence of large amounts of substrate.
However, the apparent KM is increased. In noncompetitive inhibition, substrate can bind
to the enzyme-inhibitor complex; however, the Vmax is decreased. In mixed inhibition,
both values may be altered.
Section: 8.3
Chapter 8 Mechanisms and Inhibitors 5

31. Complete the structure of the catalytic triad of chymotrypsin by drawing the proper structure of
the missing residue side chain in the box provided. Show the proper hydrogen bonding involved
in this triad.

O H O
Asp C O CH2
Ser
CH2
Ans:
His
CH2

O N H O
H N
Asp C O CH2
Ser
CH2
Section: 8.3

32. What is the challenge for a protease to facilitate hydrolysis of a peptide bond?
Ans: The peptide bond contains a carbonyl that is not very reactive; therefore, the catalytic
mechanism must employ a feature that promotes nucleophilic attack of this carbonyl
group so the peptide bond can be cleaved.
Section: 8.3

33. How can covalent modification be used to determine the mechanism of action of an enzyme?
Ans: If a particular amino acid side chain is suspected of participating in a catalytic
mechanism, covalent modification of the residue may change it enough that the enzyme
activity is altered or inhibited. However, this method is usually confirmed by other
techniques, such as site-directed mutagenesis, to rule out other possible reasons for the
loss of activity, such as conformational change.
Section: 8.3

34. What is an affinity label?

Ans: This is a substrate analog that is structurally similar to the substrate, binds to the active
site, and chemically reacts with a residue in the active site. It is used to study enzyme
structure and mechanism.
Section: 8.3

35. Why are substrate analogs used to monitor enzyme activity?

Ans: Enzyme assays must be designed so that the formation of a product is rapidly and easily
monitored. Substrates that form a colored product are easy to observe in a quantitative
manner using spectrophotometers.
Section: 8.3

36. What caused a “burst” of activity followed by a steady-state reaction when chymotrypsin was
studied by stop-flow techniques?
Ans: Chymotrypsin cleaves peptide bonds in a two-step reaction, in which the first step,
formation of the acyl enzyme intermediate, is faster than the second step, hydrolysis.
Section: 8.3
Chapter 8 Mechanisms and Inhibitors 6

37. Designing drugs to inhibit enzymes is a large part of pharmaceutical research. What are some of
the enzymatic features that would be important?
Ans: The enzyme could be inhibited by the interaction of a potential drug at the active site or at
a site that alters conformation or regulation of the enzyme. The structure of natural
substrates and activators, and their binding sites, would be useful features to study for a
new drug design. The binding affinity and specificity would be important, and standard
enzyme assays would be used to determine the effect of the inhibitors on Kcat, KM, and
Vmax.
Section: 8.2

38. What factors should an enzymologist consider when designing an enzyme assay?
Ans: Factors an enzymologist should consider when designing an enzyme assay include the pH
for optimal substrate binding and enzymatic activity, the temperature for proper catalytic
function, and the additional regulatory compounds needed to measure the enzyme’s
activity.
Section: 8.2

39. There is a key difference between an enzyme that uses a covalent catalysis mechanism and one
that uses other catalytic strategies. What is this key difference?
Ans: For each of the other strategies of catalysis, a substrate and/or a cofactor are bound to the
site by noncovalent interactions with the amino acids of the enzyme. An enzyme that uses
a covalent catalyst strategy covalently binds the substrate to one or more of the amino
acids in the active site.
Section: 8.1

40. Which of the following curves (no inhibitor, inhibitor 1, or inhibitor 2) represents the rate of
reaction versus substrate concentration for a competitive and an uncompetitive inhibitor? Draw
the double-reciprocal plot for each case.

No I
I1
V0

I2

[S]
Ans: Inhibitor 1 is a competitive inhibitor; inhibitor 2 is an uncompetitive inhibitor .
Section: 8.2

41. What is the difference between KM and KM app?

Ans: The KM is the Michaelis–Menten constant, which measures the affinity of an enzyme for
its substrate. The KM app is the altered constant in the presence of an inhibitor.
Section: 8.2 and Figures 8.10 and 8.11
Chapter 8 Mechanisms and Inhibitors 7

42. Draw and describe the reaction pathway for a noncompetitive inhibitor.
Ans: The figure should resemble the reaction pathway shown in Figure 8.9. Here, the pathway
is demonstrating that the substrate can bind irreversibly to the enzyme alone or to the
enzyme already bound to the inhibitor. The ESI complex can continue in this pathway to
an ES + I state, but may not continue to an EI + P product.
Section: 8.2

43. What are group-specific reagents?

Ans: These are compounds which will form a defined chemical reaction with specific
functional groups of an amino acid. These are used to define the active-site amino acids
of an enzyme, an example of which is diisopropylphosphofluoridate (DIPF) which
modifies a specific –OH group on a serine in chymotrypsin.
Section: 8.2

44. Bacteria that become penicillin resistant express an enzyme called -lactamase. This enzyme
hydrolyses the lactam ring on penicillin. Suggest a reason why this protein allows cells to grow
in the presence of penicillin.
Ans: Penicillin is a suicide inhibitor of the transpeptidase enzyme that modifies the serine –OH
in the active site. -lactamase reduces the concentration of the inhibitor, allowing the cell
wall of the bacteria to properly form.
Section: 8.2

45. The initial reaction kinetics of some enzymes results in a quick burst of product in a short period
of time, followed by a slower but sustained increase in product formation over time. What does
this type of kinetic response tell an enzymologist about the mechanism of the catalysis?
Ans: These two-phase reactions (a rapid-burst phase and a steady-state phase) indicate that the
reaction takes place in two steps. The first step is very quick and will achieve equilibrium
rapidly; the second step of the reaction is slower.
Section: 8.3

46. A site-directed mutagenesis converting histidine-57 of chymotrypsin to a lysine results in an

inactive enzyme even though lysine has an amino group in its side chain. Describe why the
scientist may have thought this result surprising and why it wasn’t.
Ans: The side group of histidine has two nitrogens each of which can act as either a proton
donor or an acceptor. The scientist might have thought that lysine could substitute for
histidine in the catalytic triad. However, for the triad to function properly, the substitute
would have to have the ability to partially deproteinize the –OH group of the neighboring
serine. Lysine does not have this capacity, even with a nearby aspartate.
Section: 8.3

47. How is the enzyme chymotrypsin bind and hydrolyze its substrate? How does this differ from
other proteases?
Ans: Chymotrypsin binds its substrate with a deep hydrophobic pocket, locking the substrate
into an appropriate conformation. If this binding site were altered or missing, the active
site would remain and the protein would bind many different substrates, as is the case for
some of the other proteases.
Section: 8.3
Chapter 8 Mechanisms and Inhibitors 8

48. Describe the mechanism for the proteolysis catalyzed by chymotrypsin.

Ans: This is an acid–base, covalent catalysis that generates an unstable tetrahedral-
intermediate. The intermediate is stabilized by the interactions of the oxyanion hole. The
reaction mechanism is shown on Figure 8.25.
Section: 8.3

49. You measure the initial velocity of an enzyme in the absence and presence of two inhibitors. In
each case, the inhibitor is at 10 µM. Shown in the table below is the primary data for all three
cases. Construct a Lineweaver–Burk plot for each case. Calculate the KM and Vmax for each case,
both graphically and mathematically. Determine the mechanism for each inhibitor and where
each will interact on the enzyme.
Initial Velocity (µmol/ml min)
Enzyme Enzyme Enzyme
[S] mM Alone + Inhibitor 1 + Inhibitor 2
0.33 1.65 1.05 0.79
0.50 2.13 1.43 1.02
1.00 2.99 2.22 1.43
2.00 3.72 3.08 1.79
5.00 4.00 3.80 2.00
Ans: Construct a double-reciprocal plot. From the intercept of the vertical axis, determine the
value for the intercept = 1/Vmax and the horizontal axis to determine the value for −1/KM.
Section: 8.2
Another random document with
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Volynia rose now against Volodislav. First Mystislav the Silent was
put forward, then Bailski, Leshko’s father-in-law, sent his brother,
Vsevolod, to attack the adventurer, and went himself later. Last of all
Daniel acted. After that, Leshko with Poles and men of Volynia
advanced against Volodislav. Volodislav left to his brothers the task
of defending the capital, and with hired forces hurried forth to meet
his opponents, but he was driven back and defeated. The victors
could not take Galitch, however. They fought at its walls till
exhausted, and then had to abandon the task. On the way home,
Leshko induced Bailski, now Prince of Volynia, to give two towns
near the capital to the orphans, who then moved thither from
Kamenyets, and, being near the capital, ceased not to sigh for it. “It
will come to us,” thought they. And it came earlier than they
expected.

Not Leshko, but his voevoda, Pakoslav, keen at invention, found

means to reconcile warring interests for the moment. Leshko had a
young daughter and Andrei of Hungary had a son. Leshko sent
Pakoslav to the king with this message: “Volodislav, a boyar, should
not be on a throne. Take thou my Saloméya for thy Koloman, and let
us install them in Galitch.” Pakoslav’s plan pleased Andrei. He had a
meeting with Leshko, and they [178]arranged all the details of the
marriage. The king, from the portion of Koloman, gave two cities to
Pakoslav,—Peremysl and Lubetch.

Pakoslav now offered a second good counsel: “Let the prince, out of
love for the orphans, give them Vladimir of Volynia.” Immediately
Leshko sent this message to Bailski: “Give Vladimir to Vassilko and
Daniel. If thou wilt not consent, I will take it.” Bailski would not yield,
then Leshko constrained him, and installed Roman’s sons in
Vladimir.
Volodislav, now a prisoner, was put in fetters and died in
confinement. No prince would shelter his orphans, because their
father had aspired to sovereignty.

The King and Queen of Galitch, though mere children, were crowned
straightway. Andrei, seeing that the boyars were desirous of union
with Hungary, and remembering their statement that the people
would not oppose union, if their faith and its ceremonies were
respected, now wrote to the Pope on the subject: “Let it be known to
your Holiness that the princes of Galicia, and the people there under
us, wish as king our son, Koloman, and promise union with the Most
Holy Roman Church if they may keep their own ritual. Lest delay
harm a thing so useful to us and to you, give a written command, we
beg of you, to the Archbishop of Strigonia to anoint, at the earliest,
our son, the King of Galitch.”

In the Russian chronicles it is written under the year 1214: “The

Ugrian king seated his son in Galitch; he then drove out and hunted
the clergy and bishops from the churches, and brought in his own
Latin priesthood.”

Thus Galitch was lost for a time to Russian princes and the Orthodox
clergy. In Chernigoff and Kief, people were not thinking of Galitch;
they had their own troubles. Chermny and Rurik exchanged
principalities, Chermny went to Kief and Rurik to Chernigoff. Thus
the ancient home of Oleg and his descendants passed to a
descendant of Monomach, and Chermny, the senior of Oleg’s
descendants, not only took the old capital, but threatened to drive
from Kief regions all the descendants of Monomach. He declared
that through their fault a terrible crime had been committed. “Ye
caused the death of my cousins in Galitch, and put a great shame on
us. Ye have no part in Kief regions,” asserted he. Still after that
Chermny turned to Big Nest with [179]a prayer for peace and
friendship. He begged the metropolitan to bear this request to
Vladimir. Peace was granted, and that winter Big Nest strengthened
this peace by a marriage between his second son, Yuri, and the
daughter of Chermny.

Toward the end of his life, the Prince of Vladimir had many disputes
with Novgorod, which for years had been friendly. It was most
important for Novgorod to be at peace with Vladimir, to trade with its
broad regions, and receive grain, which in Novgorod was lacking at
all times. Nothing harmed Novgorod more than a quarrel with
Vladimir, whose prince could stop grain from reaching the city and
surrounding country, and arrest Novgorod merchants wherever he
found them in his own territory. But this was not sufficient to change
the quarrelsome disposition of Novgorod, where factions fought with
one another continually. When a posadnik displeased them, they
beat him, or hurled the man from the bridge to the river. Big Nest did
not interfere with their freedom. On the contrary he apparently
commended it. “Love him who seems good to you, but execute bad
men,” said he. And the Novgorod people carried out this instruction,
even against their own adherents, the Miroshiniches, with whom they
settled in real Novgorod fashion.

Miroshka was chosen posadnik in 1187 to please Big Nest. He was

the son of Naizda, a man killed by them in the days of Andrei
Bogolyubski, for adherence to Vladimir. When Miroshka died his
descendants became famous people. Big Nest was unable for a long
time to bring about the election, as posadnik, of Miroshka’s son,
Dmitri. He could not do so till he sent his own son, Constantine, as
prince to the city. The Novgorod men then cast out the old posadnik,
and gave the office to Dmitri. This brought about a conflict with a
great citizen of Novgorod, Oleksa Bogolyubski Sbyslavich, but he
met his death very quickly.
During Constantine’s stay in Novgorod, 1205–1209, with Dmitri as
posadnik, it might be said that Big Nest ruled Novgorod as pleased
him. The execution of Oleksa is proof of this. All were astounded
when Big Nest sent this command: “Execute Oleksa without trial!”
That is, at the good judgment of Constantine. And though all men
were roused, and said on the day after the execution that the Mother
of God had dropped tears for Oleksa, the will of the prince was
accomplished. After this Dmitri [180]became so strong in his office,
and served the Grand Prince so zealously, that the four years of
Constantine’s rule passed in harmony.

When Constantine was summoned by his father to the war in

Ryazan, a large force from Novgorod marched with him under
command of Dmitri, who was greatly distinguished at the taking of
Pronsk. He was wounded severely and Big Nest detained him to be
healed in Vladimir, but he died. After his death the people in
Novgorod seized all his family property, plundered his house and the
house of his father and burned them. They sold the country places of
the son and the father, and also their servants; they took possession
of their effects and divided them. The debts due the family were left
to the prince. Still the people were not satisfied; they insisted on
punishment, and when Dmitri’s body was brought from Vladimir, they
wished to hurl it into the river. Mitrophan, the archbishop, was barely
able to stop them. When Big Nest sent his son, Sviatoslav, to the
city, the people kissed the cross in assembly not to admit any son of
Dmitri to Novgorod, and they gave his family to the prince for
imprisonment. But, though Sviatoslav received the sums due Dmitri,
and through them got much wealth, he did not obey Novgorod in this
affair. Some of the family he sent under guard to Vladimir; a few he
permitted to stay unobserved in the city.

As this uprising was directed against all adherents of Big Nest, the
Novgorod people did not escape punishment. Again he arrested
Novgorod merchants and their goods throughout the lands of
Vladimir. Great inconvenience was felt by Novgorod people, and
Oleksa’s avengers spread complaints wherever they could against
Big Nest, who, being then at the height of his power and influence,
had no effective opponents. It seemed as though no man could
refuse him obedience.

But at this juncture a prince of the smallest region in Russia,

Mystislav of Toropets, son of Mystislav the Brave, had courage to
challenge the greatness of Big Nest. On hearing how Novgorod was
treated, he offered himself to the city, a thing unheard of till that day
in Russia.

In the first years of this reign, during troubles in Novgorod, Mystislav

the Brave had inflicted defeat upon Big Nest, and now, in 1210, a
more unexpected rebuff was delivered by the son of [181]that same
prince, Mystislav the Gallant, who had grown up and strengthened in
this interval, and whose fame began with this challenge. Thus far this
young prince had appeared only in small actions, in the quarrels of
Rurik, his uncle, and in two or three raids on the Polovtsi, but on
coming to Novgorod he began a brilliant career as a hero and
defender of justice, a protector of the weak and offended, and he so
towered above other princes that he soon had no equal. Later on, he
reminded the world of his father, for he made a triumphant campaign
against the Chuds, and brought them all to obedience from border to
border of that country.

His appearance in Novgorod astonished every one by its daring, and

was crowned with incredible victory. From his small, insignificant
Toropets he came with a slender but chosen army. At Torjok he
seized Sviatoslav’s boyars and took possession of their property;
then he sent the following message to Novgorod: “I bow to Holy
Sophia; to the grave of my father, and to all men of Novgorod. I have
heard of the violence done by your princes, and I grieve for my
inheritance. Do ye wish me to be prince in your city?” The Novgorod
men were delighted and sent for him. Sviatoslav they confined in the
bishop’s palace with all his attendants, to keep him till “Lord
Novgorod” should settle with his father.

The Prince of Vladimir in anger sent against “The Gallant” a

numerous army, with his three elder sons at the head of it. But
immediately after he hesitated. He now, as on a time Dolgoruki, his
father, had done, thought proper to say when he faced an untamable
enemy: “I am old, he is young in all the passions of this world. It is
not for me, near the end of existence, to be occupied with quarrels
and bloodshed. I should be patient.” And he sent envoys to Mystislav
with this message: “Thou art my son; I am thy father. Free Sviatoslav
with his boyars, and return what thou hast taken. The merchants and
their goods will be liberated.”

Mystislav did at once all that was asked of him, and Big Nest fulfilled
his promise. Sviatoslav returned to his father, and Mystislav entered
Novgorod, rejoicing that he had passed through great peril without
bloodshed.

Big Nest was nearing the end of his earthly existence. He had
continued the task undertaken by his father and his brother to
[182]preserve and enlarge the principality of Vladimir. He had not
worked for all Russia, though he had tried to hold a share in the
Russia outside of Vladimir. During his rule, which was firm and at
times even terrible, he not only preserved unimpaired, but extended
and strengthened Vladimir. He established the beginning of a state in
the North and fixed its central region. Earlier than Big Nest, not only
in the time of his father, but also in that of Andrei, his brother, Rostoff
and Suzdal were remembered as belonging to Novgorod. Men did
not consider Vladimir or Moscow or any other place, as that Great
Russia which they were to obey, and to which they must gravitate.
Before Big Nest’s activity, Bailozersk and Galitch beyond the Volga,
and other places, if not claimed by Novgorod altogether, were
claimed at least partially. Now the Dvina country beyond the Volga
had become so connected with Vladimir that all was reconstituted.
That broad region looked on itself as Great Russia, and all men
began to regard it in that light. Lord Novgorod itself was forced to
count those lands as lost forever. Neither Rostoff nor Suzdal, from
the time of Big Nest, dared to think of their earlier primacy, the
memory of which became mingled with traditions of its ancient
connection with Novgorod. After Big Nest there could be no talk of
separation from Vladimir, for it became clear that not to Rostoff, or to
Suzdal did that Great Russia gravitate, but to Vladimir.

As his father had left Rostoff and Suzdal to his younger sons and
Vladimir to the eldest, so Big Nest, almost on the eve of his death,
gave Vladimir to his eldest son, Constantine, and left Rostoff to Yuri
his second son.

Constantine, who was in Rostoff at this time and enjoyed there great
friendship among boyars, was angry that his favorite city was given
not to him but to Yuri, and he would not abandon Rostoff for Vladimir
at the command of Big Nest. This was not his first disobedience. His
father had not forgotten the campaign of Ryazan, when Constantine
spoke against him in the presence of others. Big Nest repeated the
command. Constantine refused a second time, and sent a demand
that Rostoff should be given with Vladimir. The Grand Prince was
grieved and distressed at his son’s disobedience, and there was no
measure to his anger. As a result that took place which up to this
time had been unknown in Russia: Big Nest deprived his eldest son
of [183]seniority, and gave it to his second son, Yuri. From all the
districts and towns in Vladimir he summoned an assembly of priests,
merchants, nobles, and people, with Yoan, the bishop, at the head of
them, and in their presence gave the capital of Vladimir to Yuri,
imposing on him seniority. He commanded Yuri’s brothers to obey
him, and they kissed the cross to do so. Then the people kissed the
cross to the Grand Prince, that they would obey Yuri. From this came
endless contention in the family of Big Nest, who died shortly after.
He expired at the age of fifty-eight, Sunday, April 15, 1212, at the
hour when mass was ending in all the churches of Vladimir. They
buried him near his brother Andrei in the golden-domed cathedral,
the day following his death, as was the custom at that time.

After this began ceaseless troubles, not in Galitch, Kief, and

Chernigoff, where there was never an end to trouble, but in Vladimir,
where for thirty-seven years peace and quiet had flourished.
Deprived of seniority, Constantine did not accept the decision of his
father, but warred against Yuri and Yaroslav, who stood firmly
together. Vladimir and Sviatoslav wavered, joining now one, now the
other side. Vladimir, the youngest brother, wished Moscow as his
part, but expelled from Moscow by Yuri, he obtained his father’s
inheritance in the South,—Gorodok and Pereyaslavl. Yuri offered
Constantine peace, and even Vladimir, but asked Rostoff for himself.
Constantine would not yield; he would give Suzdal, and take
Vladimir, only if Rostoff were given him also.

Yuri freed the Ryazan and Murom princes imprisoned by his father.
Strengthened by them, he could war against his brother more
successfully. Constantine, leaving for a time his attempt on Vladimir,
continued hostile action in northern places. He seized Saligalsk, and
burned Kostroma. The whole principality was in conflict from Vologda
to Moscow. A second and a third year after the death of Big Nest this
struggle continued.

Finally, Mystislav the Gallant, their now all-powerful neighbor in

Novgorod, the main decider of wars and disputes in Russia at that
time, interfered. He had made two campaigns against the Fins near
the Baltic, and inflicted sharp punishment, but he was eager for
weighty deeds and great actions, not on distant borders, but in
Russia. His cousins, the grandsons of the “monk loving”
[184]Rostislav, turned to him for succor, and protection. Chermny, now
prince in Kief, was driving them from Dnieper regions. “The Kief
prince will not give us a part in the Russian land,” complained they.
“Come thou and help us.”

Mystislav summoned the assembly and bowed down before

Novgorod, saying: “I am going to Kief to rescue my relatives. Will ye
aid me?” “If thou go, we will follow,” was the answer. The men
chosen set out under Tverdislav, but at Smolensk the Novgorod men
had a quarrel and killed a Smolensk man; they refused thereupon to
go farther, saying: “We promised to conduct the prince hither, but to
Kief we will not go.”

Mystislav embraced the posadnik, kissed all the officers, then he

bowed to the Novgorod men, bidding Godspeed to them, and moved
forward with only his personal following and Smolensk warriors.

The Novgorod men were not pleased with themselves, and they
halted. “Lord brothers,” said Tverdislav, “what ye decide will be done
at all hazards. The question is ought we to abandon our prince at
this juncture. In their day our fathers and grandfathers marched to
suffer at Kief when their prince commanded. It is clear that we
should act in the old way.” Pleased with this speech, they turned,
and with hurried marches overtook Mystislav.

Chermny’s fate was decided at Vyshgorod. His allies were crushed,

and he fled. Two of his cousins were captured. Ingvar of Volynia,
who accompanied Mystislav, refused the Kief throne, and Roman,
son of Mystislav of Smolensk, obtained it. Vladimir, son of Rurik,
received Smolensk in addition to districts near Kief inherited from his
father. So Chermny was unable to keep his promise to avenge Igor’s
sons and expel all descendants of Monomach from Dnieper regions.
Mystislav the Gallant now besieged Chermny in Chernigoff, and
imposed peace on him. Chermny died soon after, leaving as heir his
son Michael, who later on ruled in Kief and Novgorod. His name is
still known and revered among Russians, not because he ruled, but
because he died a martyr’s death among Mongols. [185]
[Contents]
CHAPTER VIII
 DANIEL OF GALITCH

Mystislav returned to Novgorod in triumph, but the mad rage of

factions had not become weaker in the interval, and he was forced to
take strong measures. Many boyars were banished, and many were
imprisoned. Mitrophan, the archbishop famed for justice, who had
received his office not against the will of Novgorod, but who was a
friend of Big Nest, was not acceptable to Mystislav’s adherents. He
was exiled to Toropet; more than that, he was imprisoned.

During Mystislav’s absence and after his return, Vladimir partisans

were not idle. Mystislav learned that he could not rely on support in
Novgorod. He learned also that there was even a movement to expel
him. He was not the man to let any one show him the road, hence he
called the assembly on a sudden, and took farewell of it. “I have
work to do in Russia,” said he; “and ye are free as to princes.” All
were astounded on hearing this, but Mystislav, while “standing on
one foot in Novgorod, had the other in Galitch,” whither Leshko had
called him most earnestly. Leshko had been summoned more than
once to the throne in Cracow. After the crowning of Koloman, the
small boy, and Salomeya, the little maiden, their fathers endured not
long in friendship. What the king had given Leshko in Galitch, he
withdrew very quickly. That happened which happens generally with
guardians who have squandered property confided to them. The
heirs demanded account touching management.

Roman’s sons were growing rapidly. Vassilko had reached an age

when he could sit on a horse, and take part in campaigning, while
Daniel was nearing strong manhood, and gave promise of having the
powers that would make him as great as his father. He complained
loudly of Leshko, the guardian who was keeping [186]possession of
his towns on the Būg, and his inheritance beyond it. It was at this
time that King Andrei took from Leshko what he had given him in
Galitch. Seeing no profit in the fact that his daughter was queen in
“Galicia,” Leshko looked on Hungarians with envy, since they treated
Galitch as if it belonged to them, and he could not conceal his
vexation on this point. He saw that of Russian princes there was only
one who had genius in fighting; that one was Mystislav the Gallant,
hence he invited that prince to assist him.

Mystislav, through relationship with Ingvar, was a natural ally of that

branch, and not of the heirs of Roman. His preference for Ingvar was
shown by offering him the Kief throne. When Mystislav drove
Chermny from Kief and asked Ingvar to that capital, it seems that
Leshko sent him letters, and when the victor went back to Novgorod
Leshko sent envoys who offered him Galitch. When Sudislav, the
boyar who favored Hungarians, heard of Mystislav’s coming, he
preferred Daniel, and sent to him promising assistance. But
Mystislav was too quick for this movement. As he approached
Galitch, the Hungarians withdrew, and with them went Sudislav.
Mystislav entered the capital without opposition, but, with all his
good-will for Bailski, he was glad now to see Daniel, who joined him.
Instead of ill-will there was friendship between the two princes, and
Mystislav gave Anna, his daughter, to Daniel in marriage. Daniel,
who was of the oldest line among Monomach’s descendants, was
connected now with the Smolensk branch of this line, and with the
younger line also, since Mystislav’s eldest daughter had married
Yaroslav, son of Big Nest.

To all friends of Galitch Daniel’s marriage seemed of good omen, but

to ill-wishers and enemies it was hateful. Daniel grew more and more
hostile to Leshko, and more and more stubbornly demanded the
return of the Brest lands. He complained to Mystislav of his enemy in
the guise of a guardian. “Leshko,” said he, “has taken many towns of
mine, and holds a good part of my inheritance. Assist me.” “My son,”
replied Mystislav, “I cannot make war on him now, for he has shown
friendship, but thou canst find other allies.”

Daniel acted. He won back many towns, and warred against Leshko
successfully. Leshko doubted not that this winning was effected
through Mystislav’s counsel, and in view of this he changed [187]his
plans quickly. The Poles and Hungarians, from being enemies,
suddenly became friends. Leshko sent a message to Hungary that
he yielded Galitch gladly to Koloman, his son-in-law. With that Andrei
immediately allied himself with Leshko, and declared war against
Mystislav and Daniel. Leshko led in a strong army. A still greater
force came from Hungary. Mystislav wished to attack in the rear,
hence he moved toward Zbruch, and ordered his son-in-law to
defend Galitch. He summoned Bailski, also, and commanded both
allies to retire into Galitch and defend it. But Bailski left this difficult
task to Daniel, who had to bear the whole brunt of the attack of
Hungarian and Polish forces. He yielded no whit, however, and
defended the city successfully.

The allies now raised the siege and turned all their strength against
Mystislav, who was acting in their rear. Mystislav spared his scant
forces, and, while withdrawing, commanded Daniel to march out of
Galitch and join him. It was easy to give this command, but far from
easy to obey it. Mystislav himself, by deft action, slipped away
without loss, marching quickly to Kamenyets and thence to the Ros
River.

Daniel found himself now in a perilous position. The allies with united
strength, rushed at his army. It was difficult to withdraw from Galitch,
and extremely difficult to retreat along the whole course of the
Dniester, repelling the ceaseless attacks of keen enemies who
disputed each step that his men made. These enemies were much
encouraged by Mystislav’s withdrawal. Their forces were vastly
greater, and Daniel’s men lacked provisions. Night and day marches,
cold and hunger, fighting on horseback, riding without food and sleep
for nights and days in succession; all this the young warrior had to
live through in that murderous retreat down the Dniester. The glory of
this march was shared by his father’s chief boyars, who were with
him. Even they were astonished by the valor of their leader, still a
stripling. They saw with wonder how he fought entire days without
dismounting, how he rushed in pursuit of the enemy, who only saved
themselves through the swiftness of their horses.

Daniel’s men suffered greatly from lack of food. It happened that on

the festival of Saint Dmitri, when tortured with hunger, they saw on a
sudden a long line of wagons hastening forward to market. They
seized the wagons, ate abundantly, and thanked the [188]Holy Martyr
for feeding them. When at last they reached that point in the Dniester
where they had to cross, and found no bridge or ford, they despaired
of escape; but all at once they saw many merchant boats sailing
toward them from the Oleshya. On those boats all of Daniel’s men
were conveyed to the opposite bank of the river, and given
provisions.

When Daniel brought his weary troops to Podolia, Mystislav

marveled at this deed of great skill and endurance. He showed every
favor to the young hero, gave him his favorite, his very best horse,
as a present. “Go now to Vladimir, the capital of thy Volynia,” said he,
“and we will avenge this insult most surely.” And he gave assurance
that he would go to the steppes and return with Polovtsi forces. From
his youth, steppe life in tents had been pleasing to Mystislav. In
those early days he found a wife among Polovtsi maidens.

Daniel, awaiting the time of fresh action, returned to his capital, but
Mystislav did not go directly to the Polovtsi. He appeared soon after
in Novgorod. During his absence many changes had taken place.
The prince sent from Vladimir by Yuri had been replaced by one sent
from Kief by Mystislav, son of the Smolensk prince, Roman. Neither
man pleased “Lord Novgorod.” The first, alarmed at disorders,
hastened home to his father; the second found still greater trouble.
Bloody battles took place on the streets, and again a posadnik was
murdered.

Novgorod turned to Yuri a second time. “If thy son will not stay with
us, send Yaroslav, thy brother,” said they. Yaroslav seemed indeed
just the prince needed in Novgorod, and the friends of Mystislav the
Gallant might think to find in him the prince for whom they had been
seeking a long time, for was he not intimate with Mystislav, being
married to his eldest daughter? Partisans also saw in him the best of
his family.

But when Yaroslav came to Novgorod an outburst so tremendous

was taking place in the city, that no man had ever seen its like
before. The passions of the people and the wild rage of parties had
never been so violent. It turned out, too, that Yaroslav himself was of
those called “young, but early.” Men were mistaken when they
thought to find in him a son-in-law who would agree with Mystislav.
He was a genuine Vladimir prince, hence in no way inclined to
preserve the famed liberties of Novgorod. [189]He had one thought
alone: to acquire additions for Vladimir. Mystislav’s adherents
immediately conceived a deep hatred for Yaroslav, the most irascible
among all the sons of Big Nest. So acute was their feeling that, while
warming his palace, he was threatened with banishment. To make
up for this hatred, his adherents “raised their heads and stood up for
the prince like a mountain.” They advised him to go to Torjok and
rule from there, holding Novgorod with all firmness.
Yaroslav went to Torjok, and then chose his own method. When a
message was sent to him saying: “Come thou to Novgorod,” he
seized the envoys and conveyed them to Pereyaslavl Beyond the
Forest, imprisoning them on an island. Novgorod now rose as one
man against Yaroslav. Meanwhile, he sat in Torjok very quietly,
laughed at the city, and gave command to seize Novgorod
merchants and their wares on all roads in Vladimir. A multitude of
Novgorod men were arrested and imprisoned. To add to the
misfortunes of the city, the harvest that year was a failure in
Novgorod regions. Yaroslav did not let one load of wheat reach the
city. From these severe measures, there was such hunger that
parents sold their children for bread, and unclean things were eaten.
People died on the streets, and their dead bodies lay on the roads,
where hungry dogs devoured them.

Novgorod sent envoys to Yaroslav, begging him to return to the city,

but he gave no answer, and arrested the envoys. Novgorod sent a
third time. “Come to thy place; come to Holy Sophia. If thou wilt not
come, declare thy intention,” begged they. Yaroslav, as usual,
detained the envoys. The men in confinement at this time numbered
two thousand. There was “wailing and great sorrow in Novgorod.”

All at once, in the midst of these terrors, Mystislav the Gallant

appeared in the city. No man there knew whence he came. Yaroslav,
who learned of his coming, sent a detachment to arrest him, but this
detachment surrendered to Mystislav. The first thing the gallant
prince did was to seize all known partisans of Yaroslav, and put them
in irons. Then, summoning the assembly and kissing the cross
before the whole people, he said to them: “Either the men and the
lands of Great Novgorod will be freed, or I will lay down my life for
the city.” “In life or in death we are with thee!” called out the citizens
in answer. [190]
Mystislav’s first move was to send a peaceful embassy to negotiate
with Yaroslav. He selected a priest, the most famed and beloved in
the city, as a sign that negotiations were to be carried on, not with
threats, but with love, and conscientiously. He gave command to
bow down to his son-in-law, saying: “My son, free the Novgorod
merchants and men. Treat with me kindly. Leave Torjok for thy own
place.”

Yaroslav dismissed the priest without discussion. Not only did he not
free the prisoners, but those whom he held in Torjok he put in chains,
and sent to Pereyaslavl Beyond the Forest. Their property and
goods he distributed among his attendants. The moment that news
of this came to Novgorod, Mystislav commanded to sound the bell of
the assembly. The whole city came to him. “Let us go, brothers!” said
he. “Let us rescue our lands. Let us liberate our own people!”

The war from the very beginning assumed an unusual character. The
citizens of Novgorod had a single mind this time, and the prince was
at one with the people. To take vengeance for injustice was their
war-cry. Mystislav was not simply enraged against his son-in-law; he
hated in him that inborn pride of the Vladimir princes. Knowing well
that this unendurable haughtiness rested really on power, knowing
well that they had a great multitude of people behind them, and a
very large army, the strong warrior looked forward, not to a brilliant
victory in this unequal conflict, but rather to the crown of a martyr,
and prepared for the struggle with fear, but also with fortitude. He
summoned from Pskoff that brother of his whom Mystislav the Brave
on his death-bed had committed to Yuri Zaharitch, the boyar. This
brother, Vladimir, called to join him his cousin Vladimir, son of Rurik,
he who had taken the throne of Smolensk, when Mystislav, son of
Roman of Smolensk, went to rule Kief, the old capital.
Mystislav the Gallant worked untiringly. Novgorod assembled all
possible warriors, but in number they were insufficient. Mystislav
doubted not that Yaroslav and Yuri, Grand Prince of Vladimir, would
act as one man, but he counted on Constantine, who in his eyes
seemed another victim of Vladimir’s self-will and insolence. To
Constantine, and not to Yuri, belonged the throne of Vladimir, by
right of birth. To deprive a son of his God-given inheritance was a sin
in Mystislav’s eyes, and he thought that when he entered [191]the
Vladimir principality, proclaiming to Yaroslav and Yuri that he was
there to reinstate their eldest brother, Constantine would assist him.
To instate the senior son seemed to Mystislav just and proper.

Constantine enjoyed general respect; he was mild, and not haughty.

To punish Yaroslav for his cruelty to Novgorod was identified in
Mystislav’s mind with reëstablishing the old order, violated by the
whim of Big Nest. The only fault found with the son, as Mystislav
thought, was that he liked not innovations established by Andrei
Bogolyubski—Constantine defended the ancient order. Hatred for
Andrei was inherited by Mystislav, and he looked on Big Nest as he
looked on Andrei.

When the gallant prince approached Torjok, Yaroslav withdrew, and

his Novgorod friends in that place followed promptly. Vladimir
partisans came to strengthen Yaroslav, who hurried now to join Yuri,
and the brothers marched to meet the invader.

Mystislav’s campaign was swift and decisive beyond parallel. Begun

the moment Yaroslav rejected peaceful offers with insult, the season
was most inconvenient, just when roads were breaking up at Easter,
which that year fell on April 10 Old Style. Still the campaign was
ended before the roads had dried thoroughly.

On Tuesday, March 1, Mystislav moved out of Novgorod. He

permitted his warriors to seize food, but forbade them most sternly to