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Library of Congress Cataloging-in-Publication Data
Names: Costanzo, Linda S., 1947- author.
Title: Physiology / Linda S. Costanzo.
Other titles: Physiology (Elsevier)
Description: Sixth edition. | Philadelphia, PA : Elsevier, [2018] | Includes index.
Identifiers: LCCN 2017002153 | ISBN 9780323478816 (pbk.)
To Heinz Valtin and Arthur C. Guyton, who have written so well for students of physiology
Richard, Dan, Rebecca, Sheila, Elise, and Max, who make everything worthwhile
Preface
Physiology is the foundation of medical practice. A firm grasp of its principles is essential for the medical student and the practicing physician. This book is intended for students of medicine and related disciplines who are engaged in the study of physiology. It can be used either as a companion to lectures and syllabi in discipline-based curricula or as a primary source in integrated or problem-based curricula. For advanced students, the book can serve as a reference in pathophysiology courses and in clinical clerkships.
In the sixth edition of this book, as in the previous editions, the important concepts in physiology are covered at the organ system and cellular levels. Chapters 1 and 2 present the underlying principles of cellular physiology and the autonomic nervous system. Chapters 3 through 10 present the major organ systems: neurophysiology and cardiovascular, respiratory, renal, acid-base, gastrointestinal, endocrine, and reproductive physiology. The relationships between organ systems are emphasized to underscore the integrative mechanisms for homeostasis.
This edition includes the following features designed to facilitate the study of physiology:
♦ Text that is easy to read and concise: Clear headings orient the student to the organization and hierarchy of the material. Complex physiologic information is presented systematically, logically, and in a stepwise manner. When a process occurs in a specific sequence, the steps are numbered in the text and often correlate with numbers shown in a companion figure. Bullets are used to separate and highlight the features of a process. Rhetorical questions are posed throughout the text to anticipate the questions that students may be asking; by first contemplating and then answering these questions, students learn to explain difficult concepts and rationalize unexpected or paradoxical findings. Chapter summaries provide a brief overview.
♦ Tables and illustrations that can be used in concert with the text or, because they are designed to stand alone, as a review: The tables summarize, organize, and make comparisons. Examples are (1) a table that compares the gastrointestinal hormones with respect to hormone family, site of and stimuli for secretion, and hormone actions; (2) a table that compares the pathophysiologic features of disorders of Ca2+ homeostasis; and (3) a table that compares the features of the action potential in different cardiac tissues. The illustrations are clearly labeled, often with main headings, and include simple diagrams, complex diagrams with numbered steps, and flow charts.
♦ Equations and sample problems that are integrated into the text: All terms and units in equations are defined, and each equation is restated in words to place it in a physiologic context. Sample problems are followed by complete numerical solutions and explanations that guide students through the proper steps in reasoning; by following the steps provided, students acquire the skills and confidence to solve similar or related problems.
♦ Clinical physiology presented in boxes: Each box features a fictitious patient with a classic disorder. The clinical findings and proposed treatment are explained in terms of underlying physiologic principles. An integrative approach to the patient is used to emphasize the relationships between organ systems. For example, the case of type I diabetes mellitus involves a disorder not only of the endocrine system but also of the renal, acid-base, respiratory, and cardiovascular systems.
♦ Practice questions in “Challenge Yourself” sections at the end of each chapter: Practice questions, which are designed for short answers (a word, a phrase, or a numerical solution), challenge the student to apply principles and concepts in problem solving rather than to recall isolated facts. The questions are posed in varying formats and are given in random order. They will be most helpful when used as a tool after studying each chapter and without referring to the text. In that way, the student can confirm his or her understanding of the material and can determine areas of weakness. Answers are provided at the end of the book.
♦ Teaching videos on selected topics: Because students may benefit from oral explanation of complex principles, brief teaching videos on selected topics are included to complement the written text.
♦ Abbreviations and normal values presented in appendices: As students refer to and use these common abbreviations and values throughout the
book, they will find that their use becomes second nature.
This book embodies three beliefs that I hold about teaching: (1) even complex information can be transmitted clearly if the presentation is systematic, logical, and stepwise; (2) the presentation can be just as effective in print as in person; and (3) beginning medical students wish for nonreference teaching materials that are accurate and didactically strong but without the details that primarily concern experts. In essence, a book can “teach” if the teacher’s voice is present, if the material is carefully selected to include essential information, and if great care is given to logic and sequence. This text offers a down-to-earth and professional presentation written to students and for students.
I hope that the readers of this book enjoy their study of physiology. Those who learn its principles well will be rewarded throughout their professional careers!
Linda S. Costanzo
Acknowledgments
I gratefully acknowledge the contributions of Elyse O’Grady, Jennifer Ehlers, and Dan Fitzgerald at Elsevier in preparing the sixth edition of Physiology. The artist, Matthew Chansky, revised existing figures and created new figures—all of which beautifully complement the text.
Colleagues at Virginia Commonwealth University have faithfully answered my questions, especially Drs. Clive Baumgarten, Diomedes Logothetis, Roland Pittman, and Raphael Witorsch. Sincere thanks also go to the medical students worldwide who have generously written to me about their experiences with earlier editions of the book.
My husband, Richard; our children, Dan and Rebecca; our daughter-in-law, Sheila; and our grandchildren, Elise and Max, have provided enthusiastic support and unqualified love, which give the book its spirit.
CHAPTER 1 Cellular Physiology
Understanding the functions of the organ systems requires profound knowledge of basic cellular mechanisms. Although each organ system differs in its overall function, all are undergirded by a common set of physiologic principles.
The following basic principles of physiology are introduced in this chapter: body fluids, with particular emphasis on the differences in composition of intracellular fluid and extracellular fluid; creation of these concentration differences by transport processes in cell membranes; the origin of the electrical potential difference across cell membranes, particularly in excitable cells such as nerve and muscle; generation of action potentials and their propagation in excitable cells; transmission of information between cells across synapses and the role of neurotransmitters; and the mechanisms that couple the action potentials to contraction in muscle cells.
Volume and Composition of Body Fluids, 1
Characteristics of Cell Membranes, 4
Transport Across Cell Membranes, 5
Diffusion Potentials and Equilibrium Potentials, 14
Resting Membrane Potential, 18
Action Potentials, 19
Synaptic and Neuromuscular Transmission, 26
Skeletal Muscle, 34
Smooth Muscle, 40
Summary, 43
Challenge Yourself, 44
These principles of cellular physiology constitute a set of recurring and interlocking themes. Once these principles are understood, they can be applied and integrated into the function of each organ system.
VOLUME AND COMPOSITION OF BODY FLUIDS
Distribution of Water in the Body Fluid Compartments
In the human body, water constitutes a high proportion of body weight. The total amount of fluid or water is called total body water, which accounts for 50% to 70% of body weight. For example, a 70-kilogram (kg) man whose total body water is 65% of his body weight has 45.5 kg or 45.5 liters (L) of water (1 kg water ≈ 1 L water). In general, total body water correlates inversely with body fat. Thus total body water is a higher percentage of body weight when body fat is low and a lower percentage when body fat is high. Because females have a higher percentage of adipose tissue than males, they tend to have less body water. The distribution of water among body fluid compartments is described briefly in this chapter and in greater detail in Chapter 6.
Total body water is distributed between two major body fluid compartments: intracellular fluid (ICF) and extracellular fluid (ECF) (Fig. 1.1). The ICF is contained within the cells and is two-thirds of total body water; the ECF is outside the cells and is one-third of total body water. ICF and ECF are separated by the cell membranes.
ECF is further divided into two compartments: plasma and interstitial fluid. Plasma is the fluid circulating in the blood vessels and is the smaller of the two ECF
TOTAL BODY WATER
subcompartments. Interstitial fluid is the fluid that actually bathes the cells and is the larger of the two subcompartments. Plasma and interstitial fluid are separated by the capillary wall. Interstitial fluid is an ultrafiltrate of plasma, formed by filtration processes across the capillary wall. Because the capillary wall is virtually impermeable to large molecules such as plasma proteins, interstitial fluid contains little, if any, protein.
The method for estimating the volume of the body fluid compartments is presented in Chapter 6.
Composition of Body Fluid Compartments
The composition of the body fluids is not uniform. ICF and ECF have vastly different concentrations of various solutes. There are also certain predictable differences in solute concentrations between plasma and interstitial fluid that occur as a result of the exclusion of protein from interstitial fluid.
Units for Measuring Solute Concentrations
Typically, amounts of solute are expressed in moles, equivalents, or osmoles. Likewise, concentrations of solutes are expressed in moles per liter (mol/L), equivalents per liter (Eq/L), or osmoles per liter (Osm/L). In biologic solutions, concentrations of solutes are usually quite low and are expressed in millimoles per liter (mmol/L), milliequivalents per liter (mEq/L), or milliosmoles per liter (mOsm/L).
One mole is 6 × 1023 molecules of a substance. One millimole is 1/1000 or 10 3 moles. A glucose concentration of 1 mmol/L has 1 × 10 3 moles of glucose in 1 L of solution.
An equivalent is used to describe the amount of charged (ionized) solute and is the number of moles of the solute multiplied by its valence. For example, one mole of potassium chloride (KCl) in solution dissociates into one equivalent of potassium (K+) and one
equivalent of chloride (Cl ). Likewise, one mole of calcium chloride (CaCl2) in solution dissociates into two equivalents of calcium (Ca2+) and two equivalents of chloride (Cl ); accordingly, a Ca2+ concentration of 1 mmol/L corresponds to 2 mEq/L.
One osmole is the number of particles into which a solute dissociates in solution. Osmolarity is the concentration of particles in solution expressed as osmoles per liter. If a solute does not dissociate in solution (e.g., glucose), then its osmolarity is equal to its molarity. If a solute dissociates into more than one particle in solution (e.g., NaCl), then its osmolarity equals the molarity multiplied by the number of particles in solution. For example, a solution containing 1 mmol/L NaCl is 2 mOsm/L because NaCl dissociates into two particles.
pH is a logarithmic term that is used to express hydrogen (H+) concentration. Because the H+ concentration of body fluids is very low (e.g., 40 × 10 9 Eq/L in arterial blood), it is more conveniently expressed as a logarithmic term, pH. The negative sign means that pH decreases as the concentration of H+ increases, and pH increases as the concentration of H+ decreases. Thus pH H =− + log[ ] 10
SAMPLE PROBLEM. Two men, Subject A and Subject B, have disorders that cause excessive acid production in the body. The laboratory reports the acidity of Subject A’s blood in terms of [H+] and the acidity of Subject B’s blood in terms of pH. Subject A has an arterial [H+] of 65 × 10 9 Eq/L, and Subject B has an arterial pH of 7.3. Which subject has the higher concentration of H+ in his blood?
SOLUTION. To compare the acidity of the blood of each subject, convert the [H+] for Subject A to pH as follows:
(. ).7197 19
Thus Subject A has a blood pH of 7.19 computed from the [H+], and Subject B has a reported blood pH of 7.3. Subject A has a lower blood pH, reflecting a higher [H+] and a more acidic condition.
Electroneutrality of Body Fluid Compartments
Each body fluid compartment must obey the principle of macroscopic electroneutrality; that is, each
Fig. 1.1 Body fluid compartments.
compartment must have the same concentration, in mEq/L, of positive charges (cations) as of negative charges (anions). There can be no more cations than anions, or vice versa. Even when there is a potential difference across the cell membrane, charge balance still is maintained in the bulk (macroscopic) solutions. (Because potential differences are created by the separation of just a few charges adjacent to the membrane, this small separation of charges is not enough to measurably change bulk concentrations.)
Composition of Intracellular Fluid and Extracellular Fluid
The compositions of ICF and ECF are strikingly different, as shown in Table 1.1. The major cation in ECF is sodium (Na+), and the balancing anions are chloride (Cl ) and bicarbonate (HCO3 ). The major cations in ICF are potassium (K+) and magnesium (Mg2+), and the balancing anions are proteins and organic phosphates. Other notable differences in composition involve Ca2+ and pH. Typically, ICF has a very low concentration of ionized Ca2+ (≈10 7 mol/L), whereas the Ca2+ concentration in ECF is higher by approximately four orders of magnitude. ICF is more acidic (has a lower pH) than ECF. Thus substances found in high concentration in ECF are found in low concentration in ICF, and vice versa.
Remarkably, given all of the concentration differences for individual solutes, the total solute concentration (osmolarity) is the same in ICF and ECF. This equality is achieved because water flows freely across cell membranes. Any transient differences in osmolarity that occur between ICF and ECF are quickly dissipated by water movement into or out of cells to reestablish the equality.
Creation of Concentration Differences Across Cell Membranes
The differences in solute concentration across cell membranes are created and maintained by energyconsuming transport mechanisms in the cell membranes. The best known of these transport mechanisms is the Na+-K+ ATPase (Na+-K+ pump), which transports Na+ from ICF to ECF and simultaneously transports K+ from ECF to ICF. Both Na+ and K+ are transported against their respective electrochemical gradients; therefore an energy source, adenosine triphosphate (ATP), is required. The Na+-K+ ATPase is responsible for creating the large concentration gradients for Na+ and K+ that exist across cell membranes (i.e., the low intracellular Na+ concentration and the high intracellular K+ concentration).
Similarly, the intracellular Ca2+ concentration is maintained at a level much lower than the extracellular Ca2+ concentration. This concentration difference is established, in part, by a cell membrane Ca2+ ATPase that pumps Ca2+ against its electrochemical gradient. Like the Na+-K+ ATPase, the Ca2+ ATPase uses ATP as a direct energy source.
In addition to the transporters that use ATP directly, other transporters establish concentration differences across the cell membrane by utilizing the transmembrane Na+ concentration gradient (established by the Na+-K+ ATPase) as an energy source. These transporters create concentration gradients for glucose, amino acids, Ca2+, and H+ without the direct utilization of ATP.
Clearly, cell membranes have the machinery to establish large concentration gradients. However, if cell membranes were freely permeable to all solutes, these gradients would quickly dissipate. Thus it is critically important that cell membranes are not freely permeable to all substances but, rather, have selective permeabilities that maintain the concentration gradients established by energy-consuming transport processes.
Substance and Units
Ca2+, ionized (mEq/L) 2.5b 1 × 10 4
Osmolarity (mOsm/L) 290 290
aThe major anions of intracellular fluid are proteins and organic phosphates.
bThe corresponding total [Ca2+] in extracellular fluid is 5 mEq/L or 10 mg/dL.
cpH is log10 of the [H+]; pH 7.4 corresponds to [H+] of 40 × 10 9 Eq/L.
Directly or indirectly, the differences in composition between ICF and ECF underlie every important physiologic function, as the following examples illustrate: (1) The resting membrane potential of nerve and muscle critically depends on the difference in concentration of K+ across the cell membrane; (2) The upstroke of the action potential of these same excitable cells depends on the differences in Na+ concentration across the cell membrane; (3) Excitation-contraction coupling in muscle cells depends on the differences in Ca2+ concentration across the cell membrane and the membrane of the sarcoplasmic reticulum (SR); and (4) Absorption of essential nutrients depends on the transmembrane Na+ concentration gradient (e.g., glucose absorption in the small intestine or glucose reabsorption in the renal proximal tubule).
TABLE 1.1 Approximate Compositions of Extracellular and Intracellular Fluids
Concentration Differences Between Plasma and Interstitial Fluids
As previously discussed, ECF consists of two subcompartments: interstitial fluid and plasma. The most significant difference in composition between these two compartments is the presence of proteins (e.g., albumin) in the plasma compartment. Plasma proteins do not readily cross capillary walls because of their large molecular size and therefore are excluded from interstitial fluid.
The exclusion of proteins from interstitial fluid has secondary consequences. The plasma proteins are negatively charged, and this negative charge causes a redistribution of small, permeant cations and anions across the capillary wall, called a Gibbs-Donnan equilibrium The redistribution can be explained as follows: The plasma compartment contains the impermeant, negatively charged proteins. Because of the requirement for electroneutrality, the plasma compartment must have a slightly lower concentration of small anions (e.g., Cl ) and a slightly higher concentration of small cations (e.g., Na+ and K+) than that of interstitial fluid. The small concentration difference for permeant ions is expressed in the Gibbs-Donnan ratio, which gives the plasma concentration relative to the interstitial fluid concentration for anions and interstitial fluid relative to plasma for cations. For example, the Cl concentration in plasma is slightly less than the Cl concentration in interstitial fluid (due to the effect of the impermeant plasma proteins); the Gibbs-Donnan ratio for Cl is 0.95, meaning that [Cl ]plasma/[Cl ]interstitial fluid equals 0.95. For Na+, the Gibbs-Donnan ratio is also 0.95, but Na+ , being positively charged, is oriented the opposite way, and [Na+]interstitial fluid/[Na+]plasma equals 0.95. Generally, these minor differences in concentration for small cations and anions between plasma and interstitial fluid are ignored.
CHARACTERISTICS OF CELL MEMBRANES
Cell membranes are composed primarily of lipids and proteins. The lipid component consists of phospholipids, cholesterol, and glycolipids and is responsible for the high permeability of cell membranes to lipid-soluble substances such as carbon dioxide, oxygen, fatty acids, and steroid hormones. The lipid component of cell membranes is also responsible for the low permeability of cell membranes to water-soluble substances such as ions, glucose, and amino acids. The protein component of the membrane consists of transporters, enzymes, hormone receptors, cell-surface antigens, and ion and water channels.
Fig. 1.2 Orientation of phospholipid molecules at oil and water interfaces. Depicted are the orientation of phospholipid at an oil-water interface (A) and the orientation of phospholipid in a bilayer, as occurs in the cell membrane (B).
Phospholipid Component of Cell Membranes
Phospholipids consist of a phosphorylated glycerol backbone (“head”) and two fatty acid “tails” (Fig. 1.2). The glycerol backbone is hydrophilic (water soluble), and the fatty acid tails are hydrophobic (water insoluble). Thus phospholipid molecules have both hydrophilic and hydrophobic properties and are called amphipathic. At an oil-water interface (see Fig. 1.2A), molecules of phospholipids form a monolayer and orient themselves so that the glycerol backbone dissolves in the water phase and the fatty acid tails dissolve in the oil phase. In cell membranes (see Fig. 1.2B), phospholipids orient so that the lipid-soluble fatty acid tails face each other and the water-soluble glycerol heads point away from each other, dissolving in the aqueous solutions of the ICF or ECF. This orientation creates a lipid bilayer.
Protein Component of Cell Membranes
Proteins in cell membranes may be either integral or peripheral, depending on whether they span the membrane or whether they are present on only one side. The distribution of proteins in a phospholipid bilayer is illustrated in the fluid mosaic model, shown in Figure 1.3.
♦ Integral membrane proteins are embedded in, and anchored to, the cell membrane by hydrophobic interactions. To remove an integral protein from the cell membrane, its attachments to the lipid bilayer must be disrupted (e.g., by detergents). Some integral proteins are transmembrane proteins, meaning they span the lipid bilayer one or more times; thus
Intracellular fluid
Peripheral protein Integral protein
Extracellular fluid
Gated ion channel
Simple diffusion Passive; downhill No No No
Facilitated diffusion Passive; downhill Yes No No
Primary active transport Active; uphill Yes Yes; direct No
Cotransport Secondary activea Yes Yes; indirect Yes (solutes move in same direction as Na+ across cell membrane)
Countertransport Secondary activea Yes Yes; indirect Yes (solutes move in opposite direction as Na+ across cell membrane)
aNa+ is transported downhill, and one or more solutes are transported uphill.
transmembrane proteins are in contact with both ECF and ICF. Examples of transmembrane integral proteins are ligand-binding receptors (e.g., for hormones or neurotransmitters), transport proteins (e.g., Na+-K+ ATPase), pores, ion channels, cell adhesion molecules, and GTP-binding proteins (G proteins). A second category of integral proteins is embedded in the lipid bilayer of the membrane but does not span it. A third category of integral proteins is associated with membrane proteins but is not embedded in the lipid bilayer.
♦ Peripheral membrane proteins are not embedded in the membrane and are not covalently bound to cell membrane components. They are loosely attached to either the intracellular or extracellular side of the cell membrane by electrostatic interactions (e.g., with integral proteins) and can be removed with mild treatments that disrupt ionic or
hydrogen bonds. One example of a peripheral membrane protein is ankyrin, which “anchors” the cytoskeleton of red blood cells to an integral membrane transport protein, the Cl -HCO3 exchanger (also called band 3 protein).
TRANSPORT ACROSS CELL MEMBRANES
Several types of mechanisms are responsible for transport of substances across cell membranes (Table 1.2). Substances may be transported down an electrochemical gradient (downhill) or against an electrochemical gradient (uphill). Downhill transport occurs by diffusion, either simple or facilitated, and requires no input of metabolic energy. Uphill transport occurs by active transport, which may be primary or secondary. Primary and secondary active transport processes
Lipid bilayer
Fig. 1.3 Fluid mosaic model for cell membranes.
TABLE 1.2 Summary of Membrane Transport
are distinguished by their energy source. Primary active transport requires a direct input of metabolic energy; secondary active transport utilizes an indirect input of metabolic energy.
Further distinctions among transport mechanisms are based on whether the process involves a protein carrier. Simple diffusion is the only form of transport that is not carrier mediated. Facilitated diffusion, primary active transport, and secondary active transport all involve integral membrane proteins and are called carrier-mediated transport. All forms of carriermediated transport share the following three features: saturation, stereospecificity, and competition.
♦ Saturation. Saturability is based on the concept that carrier proteins have a limited number of binding sites for the solute. Figure 1.4 shows the relationship between the rate of carrier-mediated transport and solute concentration. At low solute concentrations, many binding sites are available and the rate of transport increases steeply as the concentration increases. However, at high solute concentrations, the available binding sites become scarce and the rate of transport levels off. Finally, when all of the binding sites are occupied, saturation is achieved at a point called the transport maximum, or Tm. The kinetics of carrier-mediated transport are similar to Michaelis-Menten enzyme kinetics—both involve proteins with a limited number of binding sites. (The Tm is analogous to the Vmax of enzyme kinetics.) Tm-limited glucose transport in the proximal tubule of the kidney is an example of saturable transport.
Fig. 1.5 Simple diffusion. The two solutions, A and B, are separated by a membrane, which is permeable to the solute (circles). Solution A initially contains a higher concentration of the solute than does Solution B.
♦ Stereospecificity. The binding sites for solute on the transport proteins are stereospecific. For example, the transporter for glucose in the renal proximal tubule recognizes and transports the natural isomer D-glucose, but it does not recognize or transport the unnatural isomer L-glucose. In contrast, simple diffusion does not distinguish between the two glucose isomers because no protein carrier is involved.
♦ Competition. Although the binding sites for transported solutes are quite specific, they may recognize, bind, and even transport chemically related solutes. For example, the transporter for glucose is specific for D-glucose, but it also recognizes and transports a closely related sugar, D-galactose. Therefore the presence of D-galactose inhibits the transport of D-glucose by occupying some of the binding sites and making them unavailable for glucose.
Simple Diffusion
Diffusion of Nonelectrolytes
Simple diffusion occurs as a result of the random thermal motion of molecules, as shown in Figure 1.5. Two solutions, A and B, are separated by a membrane that is permeable to the solute. The solute concentration in A is initially twice that of B. The solute molecules are in constant motion, with equal probability that a given molecule will cross the membrane to the other solution. However, because there are twice as many solute molecules in Solution A as in Solution B, there will be greater movement of molecules from A to B than from B to A. In other words, there will be net diffusion of the solute from A to B, which will continue until the solute concentrations of the two solutions become equal (although the random movement of molecules will go on forever).
Fig. 1.4 Kinetics of carrier-mediated transport. Tm, Transport maximum. Membrane
Net diffusion of the solute is called flux, or flow (J), and depends on the following variables: size of the concentration gradient, partition coefficient, diffusion coefficient, thickness of the membrane, and surface area available for diffusion.
CONCENTRATION GRADIENT (CA CB)
The concentration gradient across the membrane is the driving force for net diffusion. The larger the difference in solute concentration between Solution A and Solution B, the greater the driving force and the greater the net diffusion. It also follows that, if the concentrations in the two solutions are equal, there is no driving force and no net diffusion.
PARTITION COEFFICIENT (K)
The partition coefficient, by definition, describes the solubility of a solute in oil relative to its solubility in water. The greater the relative solubility in oil, the higher the partition coefficient and the more easily the solute can dissolve in the cell membrane’s lipid bilayer. Nonpolar solutes tend to be soluble in oil and have high values for partition coefficient, whereas polar solutes tend to be insoluble in oil and have low values for partition coefficient. The partition coefficient can be measured by adding the solute to a mixture of olive oil and water and then measuring its concentration in the oil phase relative to its concentration in the water phase. Thus K
Concentration in olive oil
Concentration in water =
DIFFUSION COEFFICIENT (D)
The diffusion coefficient depends on such characteristics as size of the solute molecule and the viscosity of the medium. It is defined by the Stokes-Einstein equation (see later). The diffusion coefficient correlates inversely with the molecular radius of the solute and the viscosity of the medium. Thus small solutes in nonviscous solutions have the largest diffusion coefficients and diffuse most readily; large solutes in viscous solutions have the smallest diffusion coefficients and diffuse least readily. Thus
THICKNESS OF THE MEMBRANE (ΔX)
The thicker the cell membrane, the greater the distance the solute must diffuse and the lower the rate of diffusion.
SURFACE AREA (A)
The greater the surface area of membrane available, the higher the rate of diffusion. For example, lipid-soluble gases such as oxygen and carbon dioxide have particularly high rates of diffusion across cell membranes. These high rates can be attributed to the large surface area for diffusion provided by the lipid component of the membrane.
To simplify the description of diffusion, several of the previously cited characteristics can be combined into a single term called permeability (P) Permeability includes the partition coefficient, the diffusion coefficient, and the membrane thickness. Thus
DDiffusion coefficient
KBoltzmann constant
TAbsolute temp
rMolecular radius
Viscosity of the medium ()
By combining several variables into permeability, the rate of net diffusion is simplified to the following expression:
JPAC C AB=−() where
JNet rate of diffusion mmol/s
PPermeability(cm/s
ASurfa = = = () ) c ce area for diffusion cm
CConcentration in Solution A( A () 2 = m mmol/L)
CConcentration in Solution Bmmol/L B = ()
SAMPLE PROBLEM. Solution A and Solution B are separated by a membrane whose permeability to urea is 2 × 10 5 cm/s and whose surface area is 1 cm2. The concentration of urea in A is 10 mg/mL, and the concentration of urea in B is 1 mg/mL. The partition coefficient for urea is 10 3, as measured in an oil-water mixture. What are the initial rate and direction of net diffusion of urea?
SOLUTION. Note that the partition coefficient is extraneous information because the value for permeability, which already includes the partition coefficient, is given. Net flux can be calculated by substituting the following values in the equation for net diffusion: Assume that 1 mL of water = 1 cm3 Thus
JPAC C AB=−()
The magnitude of net flux has been calculated as 1.8 × 10 4 mg/s. The direction of net flux can be determined intuitively because net flux will occur from the area of high concentration (Solution A) to the area of low concentration (Solution B). Net diffusion will continue until the urea concentrations of the two solutions become equal, at which point the driving force will be zero.
Diffusion of Electrolytes
Thus far, the discussion concerning diffusion has assumed that the solute is a nonelectrolyte (i.e., it is uncharged). However, if the diffusing solute is an ion or an electrolyte, there are two additional consequences of the presence of charge on the solute.
First, if there is a potential difference across the membrane, that potential difference will alter the net rate of diffusion of a charged solute. (A potential difference does not alter the rate of diffusion of a nonelectrolyte.) For example, the diffusion of K+ ions will be slowed if K+ is diffusing into an area of positive charge, and it will be accelerated if K+ is diffusing into an area of negative charge. This effect of potential difference can either add to or negate the effects of differences in concentrations, depending on the orientation of the potential difference and the charge on the diffusing ion. If the concentration gradient and the charge effect are oriented in the same direction across the membrane, they will combine; if they are oriented in opposite directions, they may cancel each other out.
Second, when a charged solute diffuses down a concentration gradient, that diffusion can itself generate a potential difference across a membrane called a diffusion potential. The concept of diffusion potential will be discussed more fully in a following section.
Facilitated Diffusion
Like simple diffusion, facilitated diffusion occurs down an electrochemical potential gradient; thus it requires no input of metabolic energy. Unlike simple diffusion, however, facilitated diffusion uses a membrane carrier and exhibits all the characteristics of carrier-mediated transport: saturation, stereospecificity, and competition. At low solute concentration, facilitated diffusion typically proceeds faster than simple diffusion (i.e., is facilitated) because of the function of the carrier. However, at higher concentrations, the carriers will become saturated and facilitated diffusion will level off.
(In contrast, simple diffusion will proceed as long as there is a concentration gradient for the solute.)
An excellent example of facilitated diffusion is the transport of D-glucose into skeletal muscle and adipose cells by the GLUT4 transporter. Glucose transport can proceed as long as the blood concentration of glucose is higher than the intracellular concentration of glucose and as long as the carriers are not saturated. Other monosaccharides such as D-galactose, 3-O-methyl glucose, and phlorizin competitively inhibit the transport of glucose because they bind to transport sites on the carrier. The competitive solute may itself be transported (e.g., D-galactose), or it may simply occupy the binding sites and prevent the attachment of glucose (e.g., phlorizin). As noted previously, the nonphysiologic stereoisomer, L-glucose, is not recognized by the carrier for facilitated diffusion and therefore is not bound or transported.
Primary Active Transport
In active transport, one or more solutes are moved against an electrochemical potential gradient (uphill). In other words, solute is moved from an area of low concentration (or low electrochemical potential) to an area of high concentration (or high electrochemical potential). Because movement of a solute uphill is work, metabolic energy in the form of ATP must be provided. In the process, ATP is hydrolyzed to adenosine diphosphate (ADP) and inorganic phosphate (Pi), releasing energy from the terminal high-energy phosphate bond of ATP. When the terminal phosphate is released, it is transferred to the transport protein, initiating a cycle of phosphorylation and dephosphorylation. When the ATP energy source is directly coupled to the transport process, it is called primary active transport. Three examples of primary active transport in physiologic systems are the Na+-K+ ATPase present in all cell membranes, the Ca2+ ATPase present in SR and endoplasmic reticulum, and the H+-K+ ATPase present in gastric parietal cells and renal α-intercalated cells.
Na+-K+ ATPase (Na+-K+ Pump)
Na+-K+ ATPase is present in the membranes of all cells. It pumps Na+ from ICF to ECF and K+ from ECF to ICF (Fig. 1.6). Each ion moves against its respective electrochemical gradient. The stoichiometry can vary but, typically, for every three Na+ ions pumped out of the cell, two K+ ions are pumped into the cell. This stoichiometry of three Na+ ions per two K+ ions means that, for each cycle of the Na+-K+ ATPase, more positive charge is pumped out of the cell than is pumped into the cell. Thus the transport process is termed electrogenic because it creates a charge separation and a potential difference. The Na+-K+ ATPase is responsible
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” The regiment proceeded to Glasgow ib. 1844. Marched to Edinburgh ib. 1846. Proceeded to Monmouthshire 40 1847. Removed to Weedon ib. 1848. Augmented to the India establishment ib. 1849. Embarked for Calcutta ib. 1853. CONCLUSION ib.
” Capture of a French Eagle by the battalion ib.
” Styled “THE EIGHTY-SEVENTH, or Prince ofWales’s Own IrishRegiment,” and authorised to bear on the regimental colour and appointments the word “Barrosa,” and an Eagle with a Wreathof Laurel, above the Harp 53
” The second battalion embarked for Tarifa 54
” Siege of Tarifa by the French 55
1812. Gallant defence of the place 58
” Authorised to bear the word “Tarifa” on the regimental colour and appointments ib.
” The battalion returned to Cadiz ib.
” Action at the bridge and fort of Puerto Largo 59
1813. Battle of Vittoria 60
” Bâton of Marshal Jourdan taken by the battalion 62
” Authorised to bear the word “Vittoria” on the regimental colour and appointments 63
1813. Actions in the Pyrenees 63
” Battle of the Nivelle 64
” Authorised to bear the word “Nivelle” on the regimental colour and appointments ib.
1814. Action near Salvatira 65
” Battle of Orthes ib.
” Authorised to bear the word “Orthes” on the regimental colour and appointments ib.
” Affair at Vic Bigorre 65
” Battle of Toulouse 66
” Authorised to bear the word “Toulouse” on the regimental colour and appointments 67
” Termination of the Peninsular War ib.
” Authorised to bear the word “Peninsula” on the regimental colour and appointments 68
” Embarkation of the battalion for Cork ib.
” Removed to Portsmouth 69
” Proceeded to Guernsey ib.
1815. Stationed in that island ib.
1816. Removed to Portsmouth, and subsequently to Colchester ib. 1817. The second battalion disbanded 74
SUCCESSION OF COLONELS OF THE EIGHTY-SEVENTH REGIMENT.
1796. Sir John Doyle, Bart., G.C.B. 75 1834. Sir Thomas Reynell, Bart., K.C.B. 83 1841. Hugh Viscount Gough, G.C.B. 89
A
P P E N D I X .
List of troops in South America in 1806-7 91
Memoir of Lieut.-General Sir Charles William Doyle, C.B., and G.C.H. 92
Memoir of Lieut.-Colonel Matthew Shawe, C.B. 95
List of battalions formed from men raised in 1803 and 1804, under the “Army ofReserve” and “Additional Force Acts” 97-100
P L A T E S .
Costume of the regiment in 1793 to face 1
Colours of the regiment 40
The French Eagle captured at the battle of Barrosa on the 5th of March 1811 50
Costume of the regiment in 1853 74
ORIGINAL UNIFORM OF THE 87TH REGIMENT 1793 FOR CANNONS MILITARY RECORDS
Madeley lith3, Wellington St . Strand.
HISTORICAL RECORD OF THE EIGHTY-SEVENTH
REGIMENT, OR
THE ROYAL IRISH FUSILIERS.
179
3.
The disturbed state of affairs on the continent of Europe in 1793, particularly in France, arising from the principles of the Revolution in that country, which threatened surrounding nations with universal anarchy, occasioned preparations to be made throughout the several countries, in order to oppose the dangerous doctrines which were then diffused under the specious terms of “Liberty and Equality.”
On the 1st of February 1793, the National Convention of France, after the decapitation of King Louis XVI. on the 21st of the previous month, declared war against Great Britain and Holland. Augmentations were immediately made to the regular army, the militia was embodied, and the British people evinced their loyalty and patriotism by forming volunteer associations, and by making every exertion for the maintenance of monarchical principles, and for the defence of those institutions which had raised their country to a high position among the nations of Europe.
Upwards of fifty regiments of infantry were authorised to be raised, on this emergency, in the several parts of Great Britain and Ireland, by officers and gentlemen possessing local influence, sixteen of which regiments, viz. from the Seventy-eighth to the Ninety-third, continue at this period on the establishment of the army.
Of the officers thus honored with the confidence of their Sovereign and his Government, Lieut.-Colonel John Doyle (afterwards General Sir John Doyle, Bart., and G.C.B.) was selected, to whom a letter of service was addressed on the 18th of September 1793, authorising him to raise a regiment, to consist of ten companies of sixty rank and file in each company. The corps was speedily completed, and was designated the EIGHTY-SEVENTH, or THE PRINCE OF WALES’S IRISH REGIMENT.
The following is a copy of the LetterofService, addressed by the Secretary-at-War to Major John Doyle, on the half-pay of the late One hundred and fifth regiment, dated
“WarOffice, “18thSeptember1793.
“SIR,
“I am commanded to acquaint you, that His Majesty approves of your raising a regiment of foot, without any allowance of levy money, to be completed within three months, upon the following terms, viz.:
“The corps is to consist of one company of Grenadiers, one of Light Infantry, and eight battalion companies. The Grenadier company is to consist of one captain, two lieutenants, three serjeants, three corporals, two drummers, two fifers, and fiftyseven private men. The Light Infantry company of one captain, two lieutenants, three serjeants, three corporals, two drummers, and fifty-seven men; and each battalion company of one captain, one lieutenant, and one ensign, three serjeants, three corporals, two drummers, and fifty-seven
private men, together with the usual staff officers, and with a serjeant-major and quartermaster-serjeant, exclusive of the serjeants above specified. The captain-lieutenant is (as usual) included in the number of lieutenants above mentioned.
“The corps is to have one major with a company, and is to be under your command as major, with a company.
“The pay of the officers is to commence from the dates of their commissions, and that of the non-commissioned officers and privates from the dates of their attestations.
“His Majesty is pleased to leave to you the nomination of the officers of the regiment; but the lieut.-colonel and major are to be taken from the list of lieut.-colonels or majors on half-pay, or the major from a captain on full pay. Six of the captains are to be taken from the half-pay, and the other captain and the captain-lieutenant from the list of captains or captainlieutenants on full pay. All the lieutenants are to be taken from the half-pay; and the gentlemen recommended for ensigns are not to be under sixteen years of age.
“No officer, however, is to be taken from the half-pay who received the difference on going upon the half-pay, nor is any officer coming from the half-pay to contribute any money towards the levy, but he may be required to raise such a quota of men as you may agree upon with him.
“The person to be recommended for quartermaster must not be proposed for any other commission.
“In case the corps should be reduced after it has been once established, the officers will be entitled to half-pay.
“No man is to be enlisted above thirty-five years of age, nor under five feet five inches high. Well-made, growing lads, between sixteen and eighteen years of age, may be taken at five feet four inches.
“The recruits are to be engaged without limitation as to the period or place of their service.
“The non-commissioned officers and privates are to be inspected by a general officer, who will reject all such as are unfit for service, or not enlisted in conformity to the terms of this letter.
“In the execution of this service, I take leave to assure you of every assistance which my office can afford.
Honorable George Napier. Nathaniel Cookman. Honorable Robert Mead. Percy Freke. Richard Thompson. Howe Hadfield.
Captain-Lieutenant James Magrath. Lieutenants.
John Thompson.
William Aug. Blakeney.
John Wilson.
Thomas Clarke.
James Henry Fitz Simon.
William Warren.
William Magrath.
Barton Lodge.
Ensigns.
Fleming Kells.
William Murray.
John Carrol. —— Walker.
Benjamin Johnson. —— Salmon.
Adjutant John L. Brock.
Surgeon— —— Hill.
Quartermaster—Wm. Thomson. Chaplain—Edw. Berwick.
The effective numbers were quickly recruited, and the regiment was so far formed as to be considered fit to be employed on active continental service. It was consequently embarked in the summer of 1794, as part of a force under Major-General the Earl Moira, and was sent to join the British army in Flanders, under the command of His Royal Highness the Duke of York. While on the march the EIGHTYSEVENTH regiment was attacked on the 15th of July 1794, at the outpost of Alost, by a strong corps of the enemy’s cavalry, which it repulsed, and for which act of bravery it received the thanks of the general officer in public orders. It is a circumstance worthy of being recorded in the regimental history, that the first individual of the regiment who was wounded, was the Lieut.-Colonel by whom it was raised. In the general orders of the Earl of Moira upon this occasion, “he expressed his admiration of the cool intrepidity with which the
EIGHTY-SEVENTH regiment repulsed an attack from the enemy’s cavalry, at the bridge of Alost, where its commander, Lieut.-Colonel Doyle, received two severe wounds, but would not quit his regiment, until the enemy had given up the attack.” The Duke of York, in his public letter, thus mentioned the affair:—
“Head-quarters,Cortyke, “15thJuly1794.
“Lord Moira speaks highly of the conduct of the officers and men of the EIGHTY-SEVENTH regiment on this occasion, particularly of Lieut.-Colonel Doyle, commanding the corps, who was severely wounded.
(Signed) “FREDERICK. ”
179 5.
In 1795 the EIGHTY-SEVENTH regiment was sent into Bergen-op-Zoom to be drilled; but soon after its arrival, the Dutch garrison revolted against the government, opened the gates, and joined the French, who entered with twenty thousand men, and made a capitulation with the EIGHTY-SEVENTH, the only British corps in the town, then commanded by Lord Dungarvan (afterwards Earl of Cork), Lieut.Colonel Doyle having been sent to England for the recovery of his wounds. The capitulation was however broken by the French, and the EIGHTY-SEVENTH were marched prisoners of war into France.
179 6.
The regiment was again filled up, and, with the Tenth foot, and some marines, was sent upon a secret expedition to the North Sea, under the command of Brigadier-General John Doyle, who had been promoted Colonel of the EIGHTY-SEVENTH, on the 3rd of May 1796, to co-operate with Admiral (the late Lord) Duncan; but, having been delayed in England until the end of September, the tempestuous
weather, usual at that season of the year in those seas, dispersed the ships and small craft by which the troops were to be landed, and put an end to the object of the expedition. The troops returned to England in the ships of war, in which they embarked under the orders of Admiral Sir Richard Bickerton.
On the 14th of October 1796, the regiment embarked for the West Indies.
Spain having united with France in hostility to Great Britain, an expedition under Lieut.-General Sir Ralph Abercromby, K.B., proceeded against the Spanish island of Trinidad, which capitulated on the 18th of February 1797. No men were killed or wounded.
Lieutenant R—— Villeneuve, of the Eighth foot, major of brigade to Brigadier-General Hompesch, was the only officer wounded, and he died of his wounds.
179 7.
After the reduction of Trinidad, the force (of which the EIGHTYSEVENTH formed part) destined for the expedition against Porto Rico, being assembled, the fleet sailed from Martinique on the 8th of April 1797, and on the 10th arrived at St. Kitt’s, where it remained for a few days. On the 17th the fleet anchored off Congrejos Point, and a landing was effected on the island of Porto Rico on the following day. The troops advanced, when it was perceived that the only point on which the town could be attacked was on the eastern side, where it was defended by the Castle and Lines of St. Christopher, to approach which it was necessary to force a way over the lagoon which formed that side of the island. This passage was strongly defended by two redoubts and gun-boats, and the enemy had destroyed the bridge connecting, in the narrowest channel, the island with the main land. After every effort the British could never sufficiently silence the fire of the enemy, who was likewise entrenched in the rear of these redoubts, to hazard forcing the passage with so small a number of troops. It was next endeavoured to bombard the town from a point to the southward of it, near to a large magazine abandoned by the
enemy. This was tried for several days without any great effect, on account of the distance. Lieut.-General Sir Ralph Abercromby, seeing that no act of vigour, or any combined operation between the sea and land services, could in any manner avail, determined to reembark the troops, which was effected during the night of the 30th of April. Four Spanish field-pieces were brought off, but not a sick or wounded soldier was left behind, and nothing of any value fell into the hands of the enemy. Sir Ralph Abercromby in his despatch alluded to the troops in the following terms: “The behaviour of the troops has been meritorious; they were patient under labour, regular and orderly in their conduct, and spirited when an opportunity to show it occurred.” The EIGHTY-SEVENTH had two rank and file killed, three wounded, and thirteen missing.
The regiment subsequently proceeded to St. Lucia, which had been captured from the French in May 1796.
179 8.
During the year 1798, the regiment remained at St. Lucia.
179 9. In December 1799, the regiment proceeded from St. Lucia to Martinique.
180 0.
The regiment was removed, in April 1800, from Martinique to Dominica.
180 1. In April 1801 the regiment embarked from Dominica for Barbadoes, and in August following proceeded to Curaçoa.
180 2.
The preliminaries of peace, which had been agreed upon between Great Britain and France in the previous year, were ratified on the 27th of March 1802; but the peace which had been thus concluded was but of short duration. Napoleon Bonaparte, who had been elected First Consul of the French Republic, showed, on several occasions, that he continued to entertain strong feelings of hostility against Great Britain.
During the year 1802, the regiment continued to be stationed at Curaçoa.
180 3.
After a few months, during which further provocations took place between the two countries, war was declared against France on the 18th of May 1803. The preparations which had been making in the French ports, the assembling of large bodies of troops on the coast, and the forming of numerous flotillas of gun-boats, justified the British government in adopting the strongest measures of defence, and in calling upon the people for their aid and services. Numerous volunteer associations were formed in all parts of the kingdom in defence of the Sovereign, the laws, and the institutions of the country. The militia was re-embodied, and the regular army was considerably augmented, under the “ArmyofReserveAct,” as shown in the Appendix, page 97.
The EIGHTY-SEVENTH regiment embarked from the island of Curaçoa for England on the 12th of January 1803, on board of the ship “De Ruyter,” which, meeting with tempestuous weather, was obliged to put into Jamaica, from whence it proceeded to Antigua, where it arrived in April 1803. The regiment proceeded to St. Kitt’s in June following.
180 4.
On the 28th of July 1804 the regiment embarked from St. Kitt’s, and on the 28th of September following it landed at Plymouth, after a service of eight years in the West Indies, having lost during that period, by the diseases incident to the climate, many officers, and between seven and eight hundred men.
On the 31st of October the regiment embarked, under the command of Lieut.-Colonel Sir Edward Butler, from Plymouth, for Guernsey, of which island Major-General Doyle had been appointed to the command and to the Lieutenant-Governorship.
The British Government, having ascertained that the King of Spain had engaged to furnish powerful aid to France, felt itself compelled to consider Spain as an enemy, and accordingly issued orders for intercepting some frigates off Cadiz, which were on their way to France with cargoes of treasure: a declaration of war was consequently issued by the Court of Madrid against Great Britain on the 12th of December 1804.
The establishment of the EIGHTY-SEVENTH regiment, which had been authorised to receive men raised in certain counties of Ireland, under the Act of Parliament, dated 14th July 1804, termed the “Additional Force Act,” was augmented by a second battalion, of which a distinct account is commenced at page 41.[6]
180 5.
On the 10th of March 1805, a detachment, consisting of twentyeight serjeants, fifteen drummers, and five hundred and twentyeight rank and file, being drafts from the levy then raising in the county of Mayo by the Honorable H. E. Browne, embarked from Ireland for Guernsey, and joined the first battalion on the 15th of April following, thus considerably augmenting the effective strength of the EIGHTY-SEVENTH regiment.
180 6.
On the 2nd of November 1805, the first battalion embarked from Guernsey, and proceeded to Portsmouth.
The first battalion of the EIGHTY-SEVENTH regiment embarked at Portsmouth on the 23rd of July 1806, and proceeded to Plymouth, where it disembarked on the 6th of September following. On the 12th of that month it embarked for South America, under the command of Lieut.-Colonel Sir Edward Butler; the effective numbers were, fifty-three serjeants, eighteen drummers, and eight hundred and five rank and file.[7]
180 7.
The first battalion arrived in the Rio de la Plata in January 1807, and disembarked on the 16th of that month near Monte Video, where it took up a position in advance, protecting the breaching batteries, it having been arranged between Brigadier-General Sir Samuel Auchmuty and Rear-Admiral Stirling to lay siege to the place. The piquets of the EIGHTY-SEVENTH, under Major Miller, were attacked by the Spaniards, who were defeated with great loss. On the 3rd of February, a practicable breach being made, the troops proceeded to storm the town, which was carried, and the citadel soon afterwards surrendered.
The EIGHTY-SEVENTH, under Lieut.-Colonel Sir Edward Butler, had three officers and sixty men killed, and three officers and eighty men wounded: total, one hundred and forty-six; strength in the field, seven hundred and eighty-eight.
Killed. Wounded.
Captain Charles Beaumont. Captain John Evans.
Lieutenant—Hugh Irwine. ” R. McCrea.
Surgeon—Wilde. Lieutenant—W. Boucher.
In the public thanks issued by Brigadier-General Sir Samuel Auchmuty, the regiment is thus mentioned:
“The EIGHTY-SEVENTH, under Lieut.-Colonel Sir Edward Butler, were equally forward; and to their credit, it must be noticed, that they were posted under the great gate, to rush into the town when it should be opened by the troops, who entered at the breach; but their ardour would not allow them to wait; they scaled the walls, and openedthemselvesapassage.
(Signed) “T. BRADFORD, “Dep.Adjutant-General.”
The EIGHTY-SEVENTH subsequently received the royal authority to bear the words “MONTE VIDEO” on the regimental colour and appointments in commemoration of the gallantry evinced in the capture of that place on the 3rd of February 1807.
On the 10th of May, Lieut.-General Whitelocke having arrived from England with reinforcements, proceeded as Commander-in-chief to prepare for the attack of Buenos Ayres. In a brilliant affair at Colonia on the 7th of June, the light company of the battalion was creditably engaged. On the 18th of June the troops embarked at Monte Video, and on the 28th of the same month landed at Ensenada da Barragon, about twenty-eight miles from Buenos Ayres, without firing a shot. Major-General John Levison Gower was the second in command to Lieut.-General Whitelocke, and the EIGHTY-SEVENTH were posted in the right brigade under Brigadier-General Sir Samuel Auchmuty.
In the assault of Buenos Ayres on the morning of the 5th of July 1807, the EIGHTY-SEVENTH were formed by wings, the right commanded by Lieut.-Colonel Sir Edward Butler, and the left by Major Miller. The orders were to pierce by the two streets to the right of the Retiro, in performing which (in company with the Thirty-eighth regiment) they suffered very severely. In the course of this service, Lieutenant William Hutchinson, in command of Captain Frederick Desbarres’s company, took two of the enemy’s guns, turned them on the Plazo del Toro, and, after a few rounds, the enemy, to the number of fifteen hundred, surrendered to him. The thanks of Sir Samuel
Auchmuty were given to Lieutenant Hutchinson for his gallant conduct upon this occasion. Serjeant Byrne also distinguished himself by his bravery. Twenty-nine pieces of artillery, with a quantity of military stores, were taken. The light company, which was detached from the regiment, was taken prisoners in the convent of St. Domingo, and remained for three days, when it was restored agreeably to the articles of the treaty.
The loss of the EIGHTY-SEVENTH on this occasion was seven officers and eighty men killed, and ten officers and three hundred and twenty men wounded: total, four hundred and seventeen; strength in the field, six hundred and forty-two; remained, two hundred and twenty-five.
Killed. Wounded.
Capt. David Considine. Major Francis Miller.
” Noblet Johnston. Capt. Alexander Rose.
” Peter Gordon. ” Frederick Desbarres.
” Henry Blake.
Lieut.—Robert Hamilton. Lieut.—James O’Brien.
” Michael Barry. ” Edward Fitzgerald. Quartermr. Wm. Buchanan. ” William Crowe. Assist.-Surgeon Buxton. ” Hen. Taylor Budd.
” Robert John Love.
Ensign Godfrey Green.
The evening before this attack, there was an order that the great coats and kits of the regiment should be left in the house which the commanding officer had occupied, under the charge of the Quartermaster; or, in his absence, of a subaltern officer, with the sick or lame men who could not march, as a guard for this baggage. The Quartermaster was employed on the general staff as an assistant engineer, and the tour of duty fell upon Lieutenant Michael Barry, of the grenadier company; but this high-spirited young man earnestly solicited, and obtained permission, to accompany his regiment, and was the first officer who fell the next day. No other subaltern being found willing to remain behind, the charge was entrusted to
Quartermaster-Serjeant William Grady. He was the first man who joined the corps on its formation, and had been distinguished for his bravery, intelligence, and trustworthiness; his guard, inefficient as it must be, mustered somewhat more than twenty men. In front of the house there was a thick orchard, with a narrow path leading to it; upon this he placed double sentries during the night, and a piquet of half his force in the day-time. It appeared that at the further end of this orchard a mounted body of the enemy was concealed; these men had been posted in advance of the town, but being unable to return, in consequence of the British troops having got between it and their position, they determined to get into the country by this narrow pass; but when they rushed out of the orchard, they were fired upon by Serjeant Grady’s sentries, and, their leader falling, they retreated into their cover, and after several ineffectual attempts to escape in that direction, the party, consisting of two officers and seventy men, well mounted and armed, surrendered to the Quartermaster-Serjeant’s small force. Having secured their arms and ammunition, he marched the prisoners to head-quarters, and delivered them up to Major-General Gower. Two hours after he received written orders from Lieut.-General Whitelocke to return the arms, &c. to the prisoners, who were released, and not to fire upon or stop any party, whether armed or not, going into or coming out of the town. At nine o’clock the next night, upwards of five hundred mounted men came out of the town and surrounded the house, the owner of which was an officer of the party, who, in addition to national hostility, was in a state of great irritation at his house having been taken from him, and, as he stated, plundered by the advanced guard of the British army. They surrounded and made prisoners Serjeant Grady and his party, who had orders not to fire upon any armed body. They were marched into the town, and thrust into loathsome dungeons. The Serjeant was a peculiar object of revenge, because he refused to accept a commission in their service, and to drill their troops. This brave and excellent soldier was subsequently rewarded for his exemplary conduct by being appointed Quartermaster to the second battalion of the regiment. After the
capture of Serjeant Grady and his party, the stores were plundered, and the baggage carried off or destroyed.
Notwithstanding the intrepidity displayed by the troops, the enterprise failed. On the morning of the 6th of July the GovernorGeneral Liniers sent a letter to Lieut.-General Whitelocke, offering to restore the prisoners taken in the action of the preceding day, and also those made with Brigadier-General Beresford, on condition that the whole of the British forces should be withdrawn from South America, which proposals were accepted. The Lieutenant-General’s conduct subsequently became the subject of inquiry by a courtmartial, and he was cashiered.
During the attack upon Buenos Ayres, a number of the Spanish and native soldiers were seen in the uniform of the EIGHTY-SEVENTH regiment; this was accounted for by a ship with the clothing for that regiment, which had been sent out from England to Monte Video, having been captured and carried into Buenos Ayres by a Spanish privateer, and the clothing had thus been distributed to the armed populace.
Subsequently to the assault on Buenos Ayres, the Commander of the Forces issued the following general order:
“BuenosAyres,8thJuly1807.
“GENERAL ORDER.
“Volunteer Peter Benson Husband, of the EIGHTY-SEVENTH regiment, is appointed Ensign in that corps, in consequence of his very gallant behaviour on the morning of the 5th instant.
(Signed) “T. BRADFORD, “Deputy-Adjutant-General.”
The EIGHTY-SEVENTH returned to Monte Video, after the cessation of hostilities, and was completed by volunteers from the different corps returning to Europe. On the 2nd of August the regiment embarked for the Cape of Good Hope; and on the 4th of September following it
landed at Simon’s Bay, and marched to Cape Town, where it formed part of the garrison.
180
8. 180 9.
During the years 1808 and 1809, the first battalion continued to be stationed at the Cape of Good Hope.
181 0.
On the 23rd of October 1810, the first battalion embarked from the Cape, having been selected to form part of an expedition designed to co-operate with troops from India, under the command of Lieut.-General John Abercromby, in the capture of the Mauritius. A landing of the troops from India had taken place a few days before the division from the Cape, under Major-General William Cockell, had arrived. Its appearance off the island was, however, particularly opportune, as the French governor had previously resolved to defend his lines before Port Louis; but when he saw the force from the Cape approach the island, he relinquished the hope of being able to make effectual resistance, and surrendered this valuable colony to the British. The battalion disembarked at Port Louis on the 1st of December, where it remained on duty, after the other regiments composing the expedition returned to their respective quarters. Captain Henry C. Streatfeild with two officers and one hundred men were embarked on board a ship of war, in advance of the expedition, and landed before the force from the Cape.
181 1 to 181 4.
The first battalion continued to form part of the garrison of the island during the four following years.
181 5.
In May 1815, the first battalion at the Mauritius was directed to hold itself in readiness for active service in India, and embarked on board of transports on the 16th of June, and landed at Fort William, in Bengal, on the 3rd of August.
The light company embarked in an Arab ship, with the flank companies of the Twelfth and Twenty-second regiments, and were carried into the Gulf of Manaar; the ship being there weather-bound, the troops were landed, with the assistance of country-boats, at Calpenterre, in Ceylon, and having remained fourteen days at Point de Galle, embarked again in the Arab ship for Calcutta, where they arrived, and rejoined the regiment on the 25th of September.
On the 1st of October 1815, the first battalion of the EIGHTY-SEVENTH regiment embarked in boats, and sailed for Berhampore, where it arrived on the 14th, and again embarked and sailed for Dinapore on the 13th of November, at which place it disembarked on the 18th of December.
181 6.
The Rajah of Nepaul having broken the terms of treaty made by him with the Honorable East India Company, the battalion marched for his territories on the 15th of January 1816, and arrived at Bullvee Camp on the 24th, where it joined the army under the command of Major-General Sir David Ochterlony, K.C.B., who commanded the forces assembled on the frontiers of Nepaul; on the 3rd of February the brigades advanced by their respective routes into Nepaul, Sir David Ochterlony remaining in company with the third and fourth brigades (to the former of which the EIGHTY-SEVENTH belonged), and marched through the forest at the foot of the Nepaul Hills on the 9th. The light company of the battalion with those of the native infantry of the brigade with two guns under the command of Lieut. John Fenton, formed the advanced guard, and had a very arduous
duty to perform, in carrying the guns through the forest, which was accomplished by the personal exertions of each individual. On the 10th, the third brigade arrived at Semul Cassa Pass, and at nine o’clock A. M. the light company of the EIGHTY-SEVENTH, commanded by Lieutenant Fenton, accompanied by Sir David Ochterlony, was drawn up the pass, a height of thirty feet, by the officers’ sashes, the brigade then about five miles from the pass; on the 19th it reached the village of Etoundah on the banks of the Rapti. The advanced guard again exerted themselves in opening a communication between the third and fourth brigades through the Cheria Ghanty Pass.
On the 27th of February it arrived at Muckwanpore, and on the 28th the brigade was ordered to take possession of the heights of Sierapore, and reconnoitre the position of the enemy. Lieutenant Thomas Lee, with a piquet of forty men of the EIGHTY-SEVENTH, and strong piquets of native infantry, under their own officers, was directed to take possession of the deserted height of Sierapore. Captain Pickersgill, acting Quartermaster-General, conducted them to their ground, where having planted them, Lieutenant Lee and twenty men of the EIGHTY-SEVENTH proceeded to reconnoitre the ground in advance: the enemy advanced to recover his position; the piquet retired, and the reconnoitering party, in danger of being cut off; had to descend a hill covered with jungle, pursued by a strong party (nearly four hundred) of the enemy, and would not have escaped but for the gallantry of two soldiers of the EIGHTY-SEVENTH, Corporal James Orr of No. 5. company, and Private Patrick Boyle of the Grenadier company, who seeing the danger of the officers, placed themselves on the pathway, and by their steadiness and fire, checked the advance of the enemy. On the officers making good their retreat, these gallant fellows retired uninjured: the corporal was promoted to the rank of serjeant at the particular desire of Major-General Sir David Ochterlony; the unfortunate habit of drinking alone prevented the promotion of the private. An action afterwards took place, in which the light company, under the command of Captain Fenton, suffered considerably. The action
commenced about noon, and ceased at six o’clock P. M., leaving the British in possession of the heights for a considerable distance from Sierapore, and of one field-piece.
In this affair Lieut.-Colonel Francis Miller commanding the battalion, and Lieutenant Fenton[8] in the command of the light company (detached in the advance), particularly distinguished themselves, and received the public thanks of Major-General Sir David Ochterlony, and also of the Marquis of Hastings, the Commander-in-chief and Governor-General of India. The regiment had ten men killed, and above thirty wounded, many of whom died; the loss of the enemy was very considerable.
The Rajah, perceiving that resistance was unavailing, sued for peace, and a treaty was concluded on the 4th of March; on the 9th of that month the battalion commenced its return to Bengal, and arrived at Amowah on the 22nd of March, where it was cantoned until the 30th of June, on which day it commenced its march to Govingunge on the river Gonduck, and embarked in boats in progress to Cawnpore; on the 17th of August, the battalion arrived at Jangemowe, within a few miles of Cawnpore, but did not disembark at the latter station, until the 10th of September. About this period the battalion became very sickly from being so long confined in boats; the hospital list amounted to about four hundred and eighty, exclusive of numbers who could not be admitted for want of room. Not less than one hundred and fifty men died in this and the following month, when the cold weather coming on, in a great measure, renovated the corps.
181 7.
On the 6th of February 1817, the regiment marched from Cawnpore towards Hattrass, which fortress the Commander-in-chief had given instructions to Major-General Marshall to besiege: the division from Cawnpore arrived before Hattrass, and joined the field army, on the 20th of February.
The pettah of the fort of Hattrass having been breached, it was resolved to storm on the evening of the 25th of February, and accordingly his Majesty’s Fourteenth regiment was appointed for that duty, and the EIGHTY-SEVENTH to cover; however, the breach being found impracticable, the troops returned to their lines, but the pettah was evacuated during the night, and taken possession of on the following morning by the British troops; batteries were immediately erected against the fort, which was heavily bombarded with shells and rockets: at length the principal magazine blew up on the 2nd of March, the explosion of which was said to be distinctly heard at Meerut, nearly two hundred miles distant.
Dya Ram, Rajah of the fortress, having determined on abandoning it, most gallantly cut his way through some of the piquets of the besieging army, and effected his escape. On the morning of the 3rd of March, the right wing of the EIGHTY-SEVENTH marched into and took possession of the fortress of Hattrass, which was reduced to a mass of ruins. On the 8th of March the regiment commenced its return to Cawnpore.
In July and August the EIGHTY-SEVENTH regiment, in Bengal, was increased by a detachment of thirteen serjeants, three drummers, and two hundred and sixty-nine rank and file, men who had been transferred on the second battalion being disbanded on the 1st of February 1817.
The regiment remained at Cawnpore until the 15th of October, when it received orders to march to Secundra, where the grand army was formed under the command of the Marquis of Hastings, against the Pindaree hordes, and having remained there until a bridge of boats was completed over the Jumna, it crossed that river on the 27th, and marched to the banks of the Sind, opposite Gualior; but the grand army being, about this time, attacked by that fatal disease, the cholera morbus, compelled the Marquis, with his troops, to retire to Erich on the Bettwah: the mortality for four or five days was very great, particularly among the natives, who died in vast numbers on the road and in the villages through which the
army passed. The EIGHTY-SEVENTH lost one subaltern (Lieutenant John Coghlan), three serjeants, and forty rank and file; total, forty-four, in three days. The army having in some measure recovered, his Lordship returned to the banks of the Sind, and took up a position at Lonaree, within twenty-one miles of Gualior, where Scindiah, with a powerful force, was ready to take the field, to support the Mahratta States, which had revolted.
181 8.
On the 14th of February 1818, the different divisions of the army were broken up, in consequence of peace being concluded, and the EIGHTY-SEVENTH returned to Cawnpore, at which station it arrived on the 26th of that month.
182 0.
On the 21st of October 1820, the regiment marched from Cawnpore for Fort William, by the new road, and arrived in that garrison on the 21st of December, a distance of six hundred and sixty miles.
182 1.
On the night of the 6th of September 1821, a very alarming fire broke out in the Honorable Company’s Dispensary, situated in Calcutta, and surrounded by many valuable houses. As soon as intelligence reached the fort, two captains and ten subalterns, with about three hundred men, immediately marched to the spot, and, by the greatest exertions, prevented the fire from spreading to the neighbouring houses. The strictness with which the armed party protected the property of the inhabitants, called forth their admiration, which was followed by the annexed letter from the Governor-General, the Marquis of Hastings.
“CouncilChamber,17thSept.1821.
“MY DEAR SIR,
“It was a great satisfaction to me, though no surprise, to learn the zealous and meritorious conduct of the detachment of the EIGHTY-SEVENTH, employed in the endeavour to stop the fire last night. As some of the men have suffered in articles of dress, to repair that damage, as well as to reward the activity of the party, the Council has directed that five hundred rupees be paid to you, which you will please to distribute according to your opinion of claims.
This mark of approbation from the Governor-General in Council, towards the party in general, was followed by one to the officers employed, each being presented with a piece of plate, accompanied by the following letter:
“CouncilChamber,18thDecember1821.
“SIR,
“The Most Noble the Governor-General in Council, being desirous to evince the sense which Government entertains of the laudable exertions of those officers of his Majesty’s EIGHTYSEVENTH regiment, who were present with the detachment sent from Fort William on the occasion of the fire at the Honorable Company’s dispensary, has commanded me to transmit to you the accompanying silver cups, with a request that you will, on the part of his Lordship in Council, present one to each of the several officers named below, who are understood to have accompanied the troops on the night of the 6th of September last.”
