AN ABSTRACT OF THE DISSERTATION OF Hyo Jin Lee for the degree of Doctor of Philosophy in Chemical Engineering presented on September 4, 2012 Title: Molecular origins of surfactant- mediated stabilization of protein Abstract approved: Joseph McGuire Nonionic surfactants are commonly used to stabilize proteins during upstream and downstream processing and drug formulation. Surfactants stabilize the proteins through two major mechanisms: (i) their preferential location at nearby interfaces, in this way precluding protein adsorption; and/or (ii) their association with protein into “complexes” that prevent proteins from interacting with surfaces as well as each other. In general, both mechanisms must be at play for effective protein stabilization against aggregation and activity loss, but selection of surfactants for protein stabilization currently is not made with benefit of any quantitative, predictive information to ensure that this requirement is met. In certain circumstances the kinetics of surface tension depression (by surfactant) in protein-surfactant mixtures has been observed to be greater than that recorded for surfactant alone at the same concentration. We compared surface tension depression by poloxamer 188 (Pluronic® F68), polysorbate 80 (PS 80), and polysorbate 20 (PS 20) in the presence and absence of lysozyme and recombinant protein, at different surfactant concentrations and temperatures. The kinetic results were interpreted with reference to a mechanism for surfactant adsorption governed by the formation of a rate-limiting structural intermediate (i.e., an “activated complex”) comprised of surfactant aggregates and protein. The presence of lysozyme was seen to increase the rate of surfactant adsorption in relation to surfactant acting alone at the same concentrations for the polysorbates while less of an effect was seen for Pluronic® F68. However, the addition of salt was observed to accelerate the surface tension depression of Pluronic® F68 in the presence of lysozyme. The addition of a more hydrophobic, surface active protein (Amgen recombinant protein) in place of lysozyme resulted in greater enhancement of surfactant adsorption than that recorded in the presence of lysozyme. A simple thermodynamic analysis indicated the presence of protein caused a reduction in ∆𝑮 for the surfactant adsorption process, with this reduction deriving entirely from a reduction in ∆𝑯. We suggest that protein accelerates the adsorption of these surfactants by disrupting their self associations, increasing the concentration of surfactant monomers near the interface. Based on these air-water tensiometry results, it is fair to expect that accelerated surfactant adsorption in the presence of protein (observed with PS 20 and PS 80) will occur with surfactants that stabilize protein mainly by their own adsorption at interfaces, and that the absence of accelerated surfactant adsorption (observed with F68) will be observed with surfactants that form stable surfactant-protein associations. Optical waveguide lightmode spectroscopy was used to test this expectation. Adsorption kinetics were recorded for surfactants (PS 20, PS 80, or F68) and protein (lysozyme or Amgen recombinant protein) at a hydrophilic solid (SiO2-TiO2) surface. Experiments were performed in sequential and competitive adsorption modes, enabling the adsorption kinetic patterns to be interpreted in a fashion revealing the dominant mode of surfactantmediated stabilization of protein in each case. Kinetic results confirmed predictions based on our earlier quantitative analysis of protein effects on surface tension depression by surfactants. In particular, PS 20 and PS 80 are able to inhibit protein adsorption only by their preferential location at the interface, and not by formation of less surface active, protein-surfactant complexes. On the other hand, F68 is able to inhibit protein adsorption by formation of protein-surfactant complexes, and not by its preferential location at the interface. ©Copyright by Hyo Jin Lee September 4, 2012 All rights reserved Molecular origins of surfactant- mediated stabilization of protein by Hyo Jin Lee A DISSERTATION submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Presented September 4, 2012 Commencement June 2013 Doctor of Philosophy dissertation of Hyo Jin Lee presented on September 4, 2012 APPROVED: Major Professor, representing Chemical Engineering Chair of the School of Chemical, Biological and Environmental Engineering Dean of the Graduate School I understand that my dissertation will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my dissertation to any reader upon request. Hyo Jin Lee, Author ACKNOWLEDGEMENTS First and foremost, I would like to express sincere appreciation and gratitude to my advisor Dr. Joseph McGuire for the continuous support from the beginning till the end of my Ph.D. study. This work would have never happened if it wasn’t for his patience, motivation, enthusiasm and immense knowledge. My sincere thanks also go to Arnold McAuley, Sekhar Kanapuram, and Dave Brems at Amgen for allowing me to work and study for my Ph.D., providing me with insightful comments. I would like to thank my husband-to-be Anthony Kim for being there for me and being so understanding for all the times I had to leave him by himself when I had to go to the lab. Last but not least, I would like to thank my family; my aunts, grandmother, brother and my parents, Youngwon Lee and Heesook Hwang for giving me infinite moral support throughout my life. CONTRIBUTION OF AUTHORS Dr.Karl Schilke was very much involved in getting all the figures and assisted in editing Chapter 2.Arnold McAuley was heavily involved in the bi-weekly discussions that contributed to this dissertation. TABLE OF CONTENTS Chapter 1- Introduction ....................................................................................................... 1 Chapter 2- Molecular origins of surfactant-mediated stabilization of protein drugs........... 3 Abstract .................................................................................................................. 4 2.1. Introduction ..................................................................................................... 5 2.2. Managing protein aggregation and adsorption loss ........................................ 6 2.2.1. Mechanisms of aggregation ............................................................ 7 2.2.1.1. Concentration-induced aggregation (Mechanism 1)…...8 2.2.1.2. Aggregation induced by conformational change (Mechanism 2)……….………………………………………...10 2.2.1.3. Aggregation induced by chemical reaction (Mechanism 3).............................................................................................…11 2.2.1.4. Nucleation-dependent aggregation (Mechanism 4)…...12 2.2.1.5. Surface-induced aggregation (Mechanism 5)………....14 2.2.2. Surfactants used in drug products ................................................ 16 2.2.2.1. Polysorbates .................................................................. 18 2.2.2.2. Poloxamers ................................................................... 19 2.2.3. Surfactant modulation of protein adsorption and aggregation ..... 21 2.2.3.1. Protein stabilization by surfactant adsorption at interfaces ................................................................................... 21 2.2.3.1.1. Expectations based on sequential and competitive adsorption experiments ............................. 22 2.2.3.1.2. On the stabilization of rFVIII by polysorbate 80 at solid-water interfaces ........................................... 24 2.2.4. Protein stabilization by surfactant-protein association ................. 25 2.2.4.1. A simple view of surfactant-protein mixtures at the airwater interface ........................................................................... 25 2.3. A testable, thermodynamic argument to guide surfactant selection ............. 28 2.3.1. Insights gained from intact and protein-depleted pulmonary surfactant ................................................................................................ 28 2.3.2. Surfactant-protein association at surfactant concentrations above the CMC ………………………………………………………………..32 2.3.3. Surfactant-protein association at surfactant concentrations below the CMC ................................................................................................. 34 2.4. Conclusions ................................................................................................. 36 2.5. References .................................................................................................... 37 TABLE OF CONTENTS (Continued) Chapter 3- Protein effects on surfactant adsorption suggest the dominant mode of surfactant-mediated stabilization of protein ...................................................................... 42 Abstract ................................................................................................................ 43 3.1. Introduction ................................................................................................... 44 3.2. Material and Methods.................................................................................... 47 3.2.1. Protein and surfactants .................................................................. 47 3.2.2.Surface tension measurements ....................................................... 47 3.3. Results ........................................................................................................... 48 3.3.1. Effect of surfactant concentration ................................................. 48 3.3.2. Effect of salt .................................................................................. 51 3.3.3. Effect of temperature and comparison to a simple model for protein-mediated acceleration of surfactant adsorption .......................... 52 3.3.4. Effect of protein ............................................................................ 58 3.4. Conclusions ................................................................................................... 60 3.5. References ..................................................................................................... 62 Chapter 4- Dominant mode of surfactant-mediated protein stabilization of protein at solid surfaces .............................................................................................................................. 64 Abstract ................................................................................................................ 65 4.1. Introduction ................................................................................................... 66 4.2. Material and Methods.................................................................................... 67 4.2.1. Materials and sample preparation.................................................. 67 4.2.2. Evaluation of adsorption kinetics .................................................. 68 4.2.2.1. Surfactant+ protein co-adsorption ................................. 69 4.2.2.2. Surfactant+protein sequential adsorption ...................... 69 4.2.2.3. Surfactant+protein co-adsorption pre-coated with surfactant .................................................................................... 70 4.3. Results and Discussion .................................................................................. 70 4.3.1. Surfactant+lysozyme co-adsorption .............................................. 70 4.3.2. Pluronic® F68+lysozyme sequential adsorption .......................... 73 4.3.3. Polysorbates+lysozyme co-adsorption pre-coated with polysorbates............................................................................................. 74 4.3.4. Surfactants+ recombinant protein co-adsorption........................... 75 4.3.5. Polysorbates+recombinant protein co-adsorption pre-coated with polysorbates ............................................................................................ 78 TABLE OF CONTENTS (Continued) 4.3.6. Effect of salt on Pluronic® F68+recombinant protein coadsorption and sequential adsorption ..................................................... 79 4.4. Conclusions ................................................................................................... 83 4.5. References ..................................................................................................... 84 Chapter 5-General conclusions ........................................................................................... 85 Bibliography ....................................................................................................................... 87 LIST OF FIGURES Figure 2.1: Schematic illustration of common mechanisms of aggregation. ..................... 9 Figure 2.2: Chemical structure of polysorbate surfactants. .............................................. 18 Figure 2.3: Chemical structure of the PEO-PPO-PEO triblock copolymer Poloxamer 188. .......................................................................................................................................... 21 Figure 2.4: Mechanisms of stabilization of proteins by surfactants, which may (a) dominate the interface and prevent protein adsorption, or (b) preferentially associate with proteins and thus prevent close approach and aggregation. ............................................. 21 Figure 2.5: Schematic illustration of surface tension depression associated with five regimes of protein-surfactant and surfactant-interface interactions. ................................ 27 Figure 2.6: Schematic of differences in kinetics and extent of surface tension depression by surfactant alone (S) or surfactant with protein (S + P), at different surfactant concentrations ([S]) or a constant protein concentration. teady-state surface tensions corresponding to S and S + P converge at sufficiently high [S]. ...................................... 28 Figure 2.7: Schematic of the rate-limiting intermediate state in the transition from a phospholipid bilayer to an interfacial monolayer. ............................................................ 29 Figure 2.8: Hypothetical mechanisms for transport of surfactant from an aggregated state to the surface. In the absence of protein (top), surfactant unimers must dissociate and migrate through the liquid to the interface. Proteins may promote formation of a structural intermediate between surfactant aggregates and adsorbed unimers, reducing the thermodynamic barriers associated with their location at the interface (bottom). ........... 33 Figure 3.1: In the absence of protein (top), surfactant monomers must dissociate and migrate through the liquid in order to adsorb. The presence of protein may facilitate aggregate disruption, leading either to an increased concentration of surfactant monomers thus enhancing adsorption (middle), or to the formation of stable, surfactant-protein complexes having little or no effect on surfactant adsorption rate (bottom). ................... 46 Figure 3.2: Adsorption isotherm (surface tension vs. time) of PS 80 and lysozyme mixture at 25 °C in 1X PBS. ............................................................................................ 49 Figure 3.3: Adsorption isotherm (surface tension vs. time) of PS 20 and lysozyme mixture at 25 °C in 1X PBS. ............................................................................................ 49 Figure 3.4: Adsorption isotherm (surface tension vs. time) of Pluronic® F68 and lysozyme mixture at 25 °C in 1X PBS. ............................................................................ 50 Figure 3.5: Effect of salt on the adsorption isotherm (surface tension vs. time) of 10 ppm Pluronic® F68 and lysozyme mixture .............................................................................. 52 LIST OF FIGURES (Continued) Figure 3.6: Effect of salt on the adsorption isotherm (surface tension vs. time) of surfactants (50 ppm Pluronic® F68, PS 80 and PS 20) and lysozyme mixture ............... 52 Figure 3.7: Adsorption isotherm (surface tension vs. time) of 5 ppm PS 80 and lysozyme mixture at 5 °C, 25 °C, and 40 °C in 1X PBS. ................................................................. 53 Figure 3.8: Adsorption isotherm (surface tension vs. time) of 5 ppm PS 20 and lysozyme mixture at 5 °C, 25 °C, and 40 °C in 1X PBS. ................................................................. 54 Figure 3.9: Adsorption isotherm (surface tension vs. time) of 5 ppm Pluronic ® F68 and lysozyme mixture at 5 °C, 25 °C, and 40 °C in 1X PBS. ................................................. 54 Figure 3.10: Arrhenius plot of adsorption during initial surface tension depression. ...... 56 Figure 3.11: Determination of thermodynamic components of the activation barrier.. ... 57 Figure 3.12: Adsorption isotherm (surface tension vs. time) of PS80 and recombinant protein mixture at 25 °C in 10 mM sodium acetate, 5% sorbitol, pH 3.5 ........................ 59 Figure 3.13: Adsorption isotherm (surface tension vs. time) of PS 20 and recombinant protein mixture at 25 °C in 10 mM sodium acetate, 5% sorbitol, pH 3.5 ........................ 60 Figure 3.14. Adsorption isotherm (surface tension vs. time) of Pluronic® F68 and recombinant protein mixture at 25 °C in 10 mM sodium acetate, 5% sorbitol, pH 3.5 ... 60 Figure 4.1: Adsorption kinetics of 90 and 900 ppm PS 80, 90 and 900 ppm PS 20, and 600 and 6000 ppm Pluronic® F68 in PBS ....................................................................... 71 Figure 4.2: Adsorption kinetics of lysozyme in PBS in the presence of 0 ppm, 900 ppm PS 80 and 900 ppm PS 20 ................................................................................................ 72 Figure 4.3: Adsorption kinetics of lysozyme in PBS in the presence of 0 ppm, 600 ppm, and 6000 ppm Pluronic® F68 .......................................................................................... 73 Figure 4.4: Adsorption kinetics of 600 ppm Pluronic® F68 followed by buffer elution and the introduction of lysozyme in PBS. ........................................................................ 74 Figure 4.5: 90 ppm polysorbates adsorption followed by 90 ppm polysorbates + lysozyme co-adsorption in PBS ........................................................................................................ 75 Figure 4.6: Adsorption kinetics of 90 and 900 ppm PS 80, 90 and 900 ppm PS 20, and 600 and 6000ppm Pluronic® F68 in 10 mM sodium acetate, 5 % sorbitol, at pH 4.0..... 77 LIST OF FIGURES (Continued) Figure 4.7: Adsorption kinetics of recombinant protein in 10 mM sodium acetate, 5% sorbitol, pH 4.0 in in the presence of 0 ppm, 900 ppm PS 80, 900 ppm PS 20, and 6000ppm Pluronic® F68 ...............................................................................................…78 Figure 4.8: 90 ppm polysorbates adsorption followed by 90 ppm polysorbates + recombinant protein co-adsorption in 10 mM sodium acetate, 5 % sorbitol, pH 4.0 ....... 79 Figure 4.9: Effect of salt concentration on the adsorption of recombinant protein in 10 mM sodium acetate, 5 % sorbitol, at pH 4.0 .................................................................... 81 Figure 4.10: Adsorption kinetics of recombinant protein in 10 mM sodium acetate, 5 % sorbitol, 250 mM sodium chloride, at pH 4.0 in the presence of 0 ppm, 600ppm, and 6000ppm Pluronic® F68 .................................................................................................. 82 Figure 4.11: Adsorption kinetics of 600ppm Pluronic® F68 followed by buffer elution and the introduction of recombinant protein in 10 mM sodium acetate, 5 % sorbitol, 250 mM sodium chloride, at pH 4.0. ....................................................................................... 83 MOLECULAR ORIGINS OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN CHAPTER 1 INTRODUCTION Aggregation phenomena in protein therapeutics have been studied and reported extensively by academia, industry and regulatory agencies. Aggregation is highly undesirable due to the profound impact on the stability of the drug product, which can result in loss of activity, unwanted immunogenic responses, and increased rate of rejection as a marketable product. Several mitigation strategies are used in the biotechnology industries to stabilize therapeutic proteins, but the addition of surfactants appears to be a general approach. Nonionic surfactants are used to protect and stabilize proteins against adsorption and surface-induced activity loss as well as aggregation in solution, either by binding to the proteins and preventing protein-protein associations, or by saturating the interface and thus minimizing adsorption and subsequent conformational changes. The following chapters encapsulate the work done to better understand the roles of surfactants and surfactant-protein complexes in modulating interfacial behavior and aggregation. Chapter 2 summarizes the roles of surfactants, proteins, and surfactantprotein complexes in modulating interfacial behavior and aggregation. These events depend on surfactant properties that may be quantified using a thermodynamic model, to provide physical/chemical direction for surfactant selection or design, and to effectively reduce aggregation and adsorption loss. A fundamental understanding of the mechanisms of aggregation and how surfactants interact with interfaces and proteins (particularly the preferential location of a surfactant at an interface, or its association with protein in 2 solution), provides guidance in selecting surfactants and excipients to reduce protein losses in a given application. Chapter 3 is a study of surface tension depression by poloxamer 188 (Pluronic® F68), polysorbate 80 (PS80), and polysorbate 20 (PS20) in the presence and absence of lysozyme and recombinant protein, at different surfactant concentrations and temperatures. The kinetic results are interpreted with reference to a mechanism for surfactant adsorption governed by the formation of a rate-limiting structural intermediate (i.e., an “activated complex”) between surfactant aggregates and protein. Chapter 4 explores adsorption of lysozyme and recombinant protein at solid/liquid interfaces in relation to poloxamer 188 (Pluronic® F68), polysorbate 80 (PS80), and polysorbate 20 (PS20) at different surfactant concentrations. Adsorption kinetics were recorded for surfactants and protein in sequential and competitive adsorption modes, enabling the adsorption kinetic patterns to be interpreted in a fashion revealing the dominant mode of surfactant-mediated stabilization of protein in each case. These results confirmed predictions based on our earlier quantitative analysis of protein effects on surface tension depression by surfactants. Finally, major conclusions made in this overall study are summarized in Chapter 5. 3 MOLECULAR ORIGINS OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN DRUGS Hyo Jin Lee,a,b Arnold McAuley,b Karl F. Schilkea and Joseph McGuirea,* a School of Chemical, Biological and Environmental Engineering, Oregon State University, Corvallis, OR 97331 b * Proces and Product Development Department Amgen Inc., Thousand Oaks, CA 91320 Corresponding Author: Joseph McGuire School of Chemical, Biological and Environmental Engineering Oregon State University 103 Gleeson Hall Corvallis, OR 97331-2702 Tel: 541-737-6306 Fax: 541-737-4600 Email: mcguirej@engr.orst.edu 4 CHAPTER 2 MOLECULAR ORIGINS OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN DRUGS Abstract Loss of activity through aggregation and surface-induced denaturation is a significant problem in the production, formulation and administration of therapeutic proteins. Surfactants are commonly used in upstream and downstream processing and drug formulation. However, the effectiveness of a surfactant strongly depends on its mechanism(s) of action and properties of the protein and interfaces. Surfactants can modulate adsorption loss and aggregation by coating interfaces and/or participating in protein-surfactant associations. Minimizing protein loss from colloidal and interfacial interaction requires a fundamental understanding of the molecular factors underlying surfactant effectiveness and mechanism. These concepts provide direction for improvements in the manufacture and finishing of therapeutic proteins. We summarize the roles of surfactants, proteins, and surfactant-protein complexes in modulating interfacial behavior and aggregation. These events depend on surfactant properties that may be quantified using a thermodynamic model, to provide physical/chemical direction for surfactant selection or design, and to effectively reduce aggregation and adsorption loss. 5 2.1. Introduction In recent years, the number of protein and peptide therapeutics reaching the marketplace has increased significantly for most major pharmaceutical and biotechnology companies. Technology has advanced greatly since the development of recombinant human insulin, the first medicine to be commercially produced by DNA cloning and manipulation [1]. Since then, rapid developments in biotechnology have enabled the commercial production of various hormones, blood factors, cytokines, and fully human monoclonal antibodies. Such therapeutic proteins are widely used to manage and treat hemophilia, cancer, diabetes, hepatitis, inflammation and other ailments. Therapeutic proteins are typically based on complex polypeptides, large molecules that are made up of a well-defined sequence of amino acids, and may be produced through a combination of chemical or biological means. Proteins adopt distinct three-dimensional structures that are usually necessary for correct function, but which are often highly sensitive to the surrounding environment and may be easily distorted or altered. Although protein drugs are generally considered to have fewer inherent side effects than traditional chemical agents, they are usually very surface-active and are more susceptible to activity loss through adsorption, structural unfolding and aggregation than small molecules. This is a substantial problem for the biopharmaceutical industry, because losses of biological activity through aggregation or surface-induced structural alteration (denaturation) are encountered throughout the production, formulation and administration of therapeutic proteins. Therefore, considerable efforts have been made to identify the causes of adsorption loss and aggregation, and to develop effective methods to minimize these detrimental effects and associated costs [2]. Proteins can often be stabilized against adsorption loss through the use of properly-chosen surfactants. Preferential location of surfactant molecules at interfaces, 6 such as the walls of glass vials or the surfaces of bubbles, may strongly inhibit adsorption of proteins and prevent their subsequent denaturation or loss. In addition, formation of surfactant-protein complexes in solution can reduce the surface activity of the proteins, thus stabilizing them toward close approach and aggregation. Aggregation can also be inhibited by a variety of non-surfactant stabilizers, which are selected based on their ability to inhibit specific molecular mechanism(s) that govern aggregation phenomena in a given system. In Section 2, below, we briefly outline the major mechanisms that contribute to aggregation and surface-induced conformational changes. Some chemical strategies used to inhibit or slow those mechanisms, including the use of surfactants, are also presented. The stabilization of proteins by surfactants, in the presence of interfaces, is discussed in Section 3. Particular emphasis is given to the mechanisms by which complexes of protein and surfactant molecules might influence thermodynamic barriers, leading to stabilization of the proteins against aggregation and adsorption loss. 2.2. Managing protein aggregation and adsorption loss Aggregation phenomena in protein therapeutics have been studied and reported extensively by academia, industry and regulatory agencies. Aggregation is highly undesirable due to the profound impact on the stability of the drug product, which can result in loss of activity, unwanted immunogenic responses, and increased rate of rejection as a marketable product [3]. Several workers have reported on the different mechanisms of aggregation, and suggest possible methods to inhibit aggregation [4-6]. Various external chemical or physical factors may be responsible. Additionally, the inherent properties of the protein (e.g. charged or hydrophobic regions) may make it unusually susceptible to aggregation. In such cases, aggregation can often be inhibited by modifying the molecular properties or by changing the external environment [7,8]. 7 Inherent properties can be effectively modified by site-directed mutagenesis [7] or chemical modification (e.g. PEGylation) [8], but such modifications may compromise the biological activity of the protein. Thus, the simplest and most common method of inhibiting aggregation is to change the nature of the environment surrounding the protein by adjusting solution conditions such as pH, or by adding stabilizers/excipients. By carefully examining the mechanism(s) responsible for the aggregation, we can identify specific changes or stabilizing molecules that will effectively inhibit that mechanism via a molecular-level interaction, and thus enhance the stability of the formulation. 2.2.1 Mechanisms of aggregation Protein aggregation occurs under different stress conditions, and produces unwanted and detrimental effects on a therapeutic drug product. Aggregation occurs through several different major mechanisms and pathways (Figure 2.1), discussed in detail in the examples below. Although a particular mechanism may identify an aggregation pathway for a particular protein, it may not be relevant for another protein. Also, more than one mechanism or pathway may be simultaneously responsible for destabilizing a protein formulation [5]. A fundamental understanding of the mechanisms of aggregation is not only valuable in identifying the cause of a problem, but also helpful in suggesting methods to suppress aggregation to an acceptable level. It may be noted that aggregation may or may not lead to precipitation or insoluble aggregates 2.2.1.1. Concentration-induced aggregation (Mechanism 1) Because proteins tend to be surface-active due to their polymeric structure and amphipathic nature [9], they can form reversible aggregates especially in high concentration formulations. Mechanism 1 begins with an association of native 8 monomers into an initially-reversible complex. As protein concentration increases or time passes, the protein complex may become irreversible and lead to aggregation (Figure 2.1a). Formation of intermolecular disulfide linkages (possibly through disulfide interchange reactions) is one cause of this irreversibility [5]. IgG antibodies have been observed to form reversible soluble aggregates in high concentration solutions. Electrostatic interactions and hydrogen bonds contribute to the self-association of IgG molecules. Hydrophobic patches in the Fc region of IgG are considered to be a major factor in inducing aggregation at higher concentrations [10]. 9 1 Higher oligomers (potentially irreversible) or Native protein Reversible oligomerization of native protein Higher oligomers (typically irreversible) 2 Native protein Conformational change / unfolding Oligomerization of non-native protein Higher oligomers (potentially irreversible) 3 Native protein Chemicallymodified protein Oligomerization of modified protein; potential recruitment of native proteins Visible particulates or precipitation 4 Native protein Critical nucleus Addition of native proteins onto nucleation sites, with partial unfolding Visible aggregates, adsorption loss or precipitation 5 Native protein Container walls / air-liquid interface Adsorption and surface-induced partial unfolding Aggregation of altered protein (as in Mechanism 2) Figure 2.1: Schematic illustration of common mechanisms of aggregation. Multiple mechanisms may be at work in any given system. Adapted with permission from [5]. Copyright © 2009 Bentham Science Publishers. Human interleukin-1 receptor anatagonist (IL-1ra) is part of the IL1/Fibroblast Growth Factor family of proteins with a predominantly β-strand secondary structure. It self-associates and aggregates without changes to secondary structure at high concentrations and elevated temperatures. This self-association induced aggregation was attributed to a positively-charged Lys96 residue in the IL-1ra molecule. Aggregation 10 affinity was dependent on the buffer ionic strength and the type of anion. Phosphate anion was found to inhibit aggregation more weakly than citrate or pyrophosphate at pH 6.5. Proteolytic removal of an unstructured N-terminal region containing another lysine residue also substantially reduced the rate of self-association. It was proposed that the anions compete for cationic sites on the protein, preventing the formation of cation-π interactions between protein molecules. Based on pK values at 25 °C, citrate and pyrophosphate anions would have 2 to 3 times more ionized groups than phosphate at pH 6.5. The relative affinity of the anion binding to the cationic site (and hence, inhibition of aggregation) was correlated with the number of ionizable groups at a given pH [7]. Insulin aggregation is generally considered reversible at room temperature near its isoelectric point. This can be attributed to electrostatic interactions due to the marked charge anisotropy of the polypeptide. Addition of heparin, a highly charged polyanion, prevented aggregation at pH higher than 6 by binding to the positive domains of insulin to form heparin-insulin complexes. Heparin was also able to dissociate aggregate particles of insulin, indicating that the association was largely charge-based [11]. 2.2.1.2. Aggregation induced by conformational change (Mechanism 2) Another very common form of aggregation occurs when non-native states of the protein have a higher affinity with each other than the native state. In Mechanism 2, proteins aggregate after they go through a conformational change or partial unfolding (denaturation), which is the rate-limiting step (Figure 2.1b). Interactions between the denatured proteins are typically driven by hydrophobic associations, and are usually strong enough to be practically irreversible. External factors like heat and shear 11 can induce the protein into a non-native state, as is commonly observed with the proteins in egg whites. Stability of interferon-tau (INF-tau), which is a novel type 1 interferon, depended on the type of buffering agent, even when the pH and ionic strength were kept constant. At pH 7.0, INF-tau in Tris and phosphate buffers at elevated temperatures was observed to aggregate, with a substantial loss of tertiary structure and slightly expanded non-native conformation. However, samples containing free histidine as a buffering agent suppressed thermally-induced aggregation, and little loss of tertiary structure was observed at the same pH. Detectable binding was observed only for histidine, and only to the native conformation. Histidine had little effect on protein-protein repulsion, suggesting that colloidal stabilization was unimportant. Thermodynamic stabilization was achieved by binding of histidine to a specific ligand in the native state of INF-tau and thus, maintains its native state and suppresses aggregation [12]. While normally a stable drug product, samples of recombinant human granulocyte colony stimulating factor (rhGCSF) were observed to aggregate under physiological conditions [13]. Added sucrose was able to stabilize rhGCSF, and inhibited aggregation under stressed conditions. The thermodynamic stability of rhGCSF increased with the addition of sucrose, which is preferentially excluded from the surface of the protein [14, 15]. In this system, sucrose acted as a stabilizer by shifting the equilibrium to favor the native compact species rather than the structurally expanded species. 2.2.1.3. Aggregation induced by chemical reaction (Mechanism 3) Mechanism 3 is similar to Mechanism 2, but the conformational change is caused by chemical modification or degradations such as oxidation, deamidation, or disulfide scrambling (Figure 2.1c). Chemical changes may profoundly alter protein 12 properties such as solvent accessibility of hydrophobic patches, reduction in electrostatic repulsion due to modification of charged residues, or disruptions of the native structure that trigger unfolding. It is important to note that chemically different species are not necessarily degradation products, but may be formed during normal production of a drug. Truncated peptides or under-glycosylated glycoproteins may be more susceptible to aggregation than their correctly-formed counterparts [5]. The stability of a basic leucine zipper (bZIP) domain of activating transcription factor 5, which consists of a single α-helix and a single cysteine residue, was observed to be dependent on intermolecular disulfide bonds which stabilize the native structure. Reduction of the disulfide bond resulted in the unfolding of the peptide and exposed hydrophobic regions, which resulted in aggregation of the protein [16]. A detailed structural characterization of the effects of methionine oxidation on the stability of the human IgG Fc region was studied. Oxidation of methionine can generate a repulsive interaction between the side chains of methioninine residues in the CH2 and CH3 domains, and thus disrupt the native structure. Although Met residues in both CH2 and CH3 domains were affected by oxidation, more structural perturbations were observed in the CH2 domain. Therefore, an increase in aggregation would be likely due to the structural instability of the CH2 domain of the Fc region. Since aggregation was observed only for highly oxidized proteins, addition of excipients such as methionine or sodium thiosulfate that acts as oxygen scavengers would be sufficient in preventing methionine oxidation that can cause aggregation [17]. 2.2.1.4. Nucleation-dependent aggregation (Mechanism 4) In contrast to the three previous mechanisms, which are based on interactions between individual protein molecules, protein aggregation can also be 13 attributed to nucleation-dependent processes. Mechanism 4 describes an aggregation process that is initiated when a “critical nucleus” is formed in solution, and native proteins are recruited, and often partially unfolded, to form aggregated species (Figure 2.1d). The process is not unlike the growth of a large crystal from a supersaturated solution after addition of a seed crystal. In this case, the “seed” is a microscopic aggregated particle of a denatured or otherwise non-native conformation. A “lag phase” (often weeks or months) is characteristic of this mechanism. During this lag phase (which may vary considerably from sample to sample), the seed nucleus grows, but no particles or precipitation can be observed. After the formation of a critical nucleus, the aggregation progresses rapidly, with the relatively sudden formation of visible aggregates or precipitates in solution. These nuclei may be denatured proteins, or solid contaminants (e.g. particles of silica from vials or metal from pumps) [5]. An excellent example of this nucleation-dependent mechanism is the 10residue peptide of human amylin, which is used as a model system to study self-assembly of amyloid fibril proteins. Amylin was observed to aggregate in response to low levels of asparagine deamidation, such as might be found as impurities in synthetic amyloid peptides. Seeding solutions of the native peptide with small amounts of deamidated peptide resulted in rapid aggregation to form characteristic amyloid structures. Additionally, when the affected side chain of the deamidated peptide is deprotonated and negatively charged (at physiological pH), electrostatic interactions with the positivelycharged N-terminus of another amylin peptide induce the propagation of the aggregation event. A relevant point is that phosphate anion is known to promote deamidation of Asp/Gln residues [18]. Tungsten contamination from a needle tip was observed to induce significant protein aggregation in pre-filled syringes. Tungsten microparticles become 14 soluble at lower pH, forming tungsten polyanions which are able to precipitate a monoclonal antibody (mAb) within seconds. The tungsten polyanions bind to the proteins, reducing the net charge and screen the electrostatic repulsions between the native monomers to induce precipitation. However, at pH 6.0 and higher, tungsten polyanions do not form and aggregation was not observed. The authors caution that the small number of tungsten particles required to induce precipitation of the antibodies, combined with poor mixing in the needle, precluded defining an acceptable syringe volume for a given protein [19]. Silicone oil, a common lubricant in pharmaceutical applications, has also been implicated in aggregation of monoclonal antibodies in pre-filled syringes. Although silicone oil alone did not induce aggregation, a synergistic effect producing substantial aggregation was observed when samples were agitated in the presence of silicone oil. Perturbation of the monomeric state of the protein by a combination of air-water and oilwater interfacial stresses was implicated in the aggregation. Complete inhibition of silicone oil-induced protein loss was observed when the nonionic surfactant polysorbate 20 was added. Polysorbate 20 is known to compete with protein molecules at air-water and oil-water interfaces (model hydrophobic systems), where it prevents the structural perturbations and subsequent aggregation of the protein molecules at the unprotected interfaces [20]. 2.2.1.5. Surface-induced aggregation (Mechanism 5) Finally, Mechanism 5 describes a surface-induced aggregation process, in which native proteins first adsorb to an interface, after which they undergo conformational changes or partial unfolding (Figure 2.1e). The resulting non-native conformation then serves as a starting point for aggregation in solution or directly on the 15 surface, as described in Mechanism 2 (above). While the previous mechanisms have dealt with proteins in solution, this “heterogeneous” mechanism requires the presence of an interface (typically air-water or solid-water). Protein binding at the air-water interface can be attributed to hydrophobic interactions, while electrostatic interactions often contribute at the solid-liquid interface. Nonionic surfactants are used in this case to protect and stabilize proteins against surface activity loss and/or surface-induced aggregation, either by binding to the proteins and preventing protein-protein associations, or by saturating the interface and thus minimizing adsorption and subsequent conformational changes. These effects will be discussed in detail in Section 3, below. The nonionic surfactants Tween 20® and Tween 80® were seen to protect albutropin, a recombinant human growth hormone–albumin fusion protein, against agitation-induced aggregation in solution [21]. Although the binding affinity between the protein and Tween® was different for different Tween® formulations, aggregation was completely inhibited by the surfactant binding directly to the protein, at concentrations well below the critical micelle concentration (CMC). The surfactants increased the protein conformational stability by increasing the free energy of unfolding associated with denaturation/aggregation. Joshi et al. investigated the stabilization of non-agitated samples of human recombinant Factor VIII (rFVIII) against aggregation in the presence of polysorbate 80. Association of the rFVIII with the surfactant in solution provided an effective steric barrier to aggregation, although shear fields were found to interfere with the stability of the polysorbate 80-rFVIII association. At high concentrations of polysorbate 80, however, the enhanced stabilization of agitated rFVIII was attributed to rapid and preferential adsorption of polysorbate 80 at nascent air-water interfaces [22]. 16 The extent of aggregation also depends upon the surface chemistry of the container in which the protein is stressed. For example, more aggregation was observed in Teflon®-like containers than in glass containers after a freeze-thaw cycle [23]. In another study, rFVIII was adsorbed on colloidal particles with hydrophilic or hydrophobic surfaces and net positive or negative surface charge densities. Hydrophilic surfaces exhibited relatively high rFVIII adsorption, but did not induce large changes in structure or biological activity. In contrast, exposure to hydrophobic nanoparticles caused substantial changes in tertiary structure and reduced the biological activity (as measured by activated partial thromboplastin time) of rFVIII. High surfactant concentrations, however, reduced these surface-induced effects due to competitive hindrance of rFVIII adsorption at the surfactant-coated surface [24]. 2.2.2. Surfactants used in drug products Surfactants are amphipathic, surface-active molecules that readily adsorb at interfaces. Although literally thousands of different surfactants are commercially available, all generally consist of a hydrophilic “head”, which can be ionic or a highly polar polymer, and a hydrophobic “tail”, often a long-chain aliphatic hydrocarbon group. Surfactants can be classified as anionic, cationic, nonionic and amphoteric based upon the nature of the hydrophilic “head”. An excellent example of a widely-used anionic surfactant is sodium dodecyl sulfate (SDS); the dodecyl (C12) tail is hydrophobic, while the sulfate group is highly polar. Surfactants have wide-spread applications in industry as emulsifiers, foaming agents, wetting agents, dispersants and detergents. The pharmaceutical and biotechnology industries primarily use nonionic surfactants for a variety of applications (including stabilization of protein therapeutics), because these surfactants exhibit low toxicity and low sensitivity to the presence of electrolytes. 17 At low concentrations, surfactants will adsorb to all available interfaces, replacing the higher energy molecules, and lowering the overall interfacial free energy of the system. However, as more surfactant molecules are introduced, eventually the interfaces become saturated. At this point, energy reduction is achieved by formation of micelles or other aggregated states, in which the hydrophobic “tails” are in the center and away from the surrounding water. The critical micelle concentration (CMC) is defined as the bulk concentration of surfactant at which micellization begins to occur, and is an important fundamental property of a surfactant. However, the CMC does not completely describe the surfactant’s effect in a protein mixture. If the surfactant has a high affinity for a surface, then surfactant concentrations near the CMC will tend to stabilize protein against surface-induced denaturation, even when no specific binding of the surfactant to the protein is observed. In contrast, if the surfactant stabilizes proteins by directly binding to them, the effective surfactant concentration is related to the ratio of surfactant to protein, rather than the CMC [25]. Equilibrium air-water interfacial tensiometry measurements of surfactant solutions at various concentrations are commonly used to estimate the CMC. In this approach, the CMC is determined as the bulk surfactant concentration beyond which the equilibrium interfacial tension is independent of surfactant concentration (i.e. adding more surfactant has no effect on the interfacial tension). This approach is often applied to identify the apparent CMC of a surfactant in protein-surfactant mixtures as well. In either case, it is implicitly assumed that when the CMC of the surfactant is met, the steady-state interfacial tension is governed entirely by the surfactant at the interface, and independent of other factors. It is important to note that experimental measures of the CMC are generally not sharp transitions, and are strongly dependent upon factors such as ionic 18 strength and temperature. Thus, literature values for a given surfactant may vary widely between reports [26]. 2.2.2.1 Polysorbates O O O w OH O O x HO O O z OH y R (Aliphatic) R= (CH2)11 (PS-20) (CH2)7CH=CH(CH2)8 (PS-80) w + x + y + z = 20 Figure 2.2: Chemical structure of polysorbate surfactants. The aliphatic (hydrophobic) tails polysorbate 20 and 80 vary in length and degree of unsaturation, while the PEO content remains constant. Polysorbates have a common structure consisting of a sorbitan ring with poly(ethylene oxide) at the hydroxyl positions, and differ only in the structures of the fatty acid side chains (Figure 2.2). Differences in the length and unsaturation of the fatty acid side chain structures cause the binding affinities of polysorbates with proteins to differ [27]. The most commonly-used nonionic surfactants are polysorbate 20 (polyoxyethylene sorbitan monolaureate), sold commercially as Tween 20®, and polysorbate 80 (polyoxyethylene sorbitan monooleate, or Tween 80®). Polysorbate 80 is considerably more surface-active and has a lower CMC than polysorbate 20, because it has a longer and monounsaturated aliphatic chain [28]. Polysorbate 80 also exhibits a weaker interaction with human serum albumin than polysorbate 20, again due to its longer fatty acid chain [21, 27]. Differences in the binding affinity and interaction of polysorbate 20 and 80 with darbepoetin alfa have been reported. Polysorbate 80 binds to darbepoetin alfa with minimal effect on its tertiary structure, while binding of polysorbate 20 binding caused partial unfolding of the protein [29]. As mentioned above, polysorbates are widely reported to suppress aggregation upon agitation, shaking, freeze-drying and freeze-thawing processes, and can 19 substantially prevent protein adsorption at solid surfaces [21, 24, 30-33]. However, the effectiveness appears to depend on the particular stress involved: in one study, stirring of an antibody suspension was found to be more stressful than shaking, despite the renewal of air-water interfaces during shaking. Polysorbate 20 at concentrations above 0.0025% (w/v) inhibited aggregation during shaking. However, at lower polysorbate concentrations, the protein was destabilized by shaking, and much higher surfactant concentrations were required to stabilize stirred suspensions [34]. The polysorbates are susceptible to autoxidation at moderate temperatures, primarily by radical reactions at the PEO and unsaturation sites of the olefinic moieties, and hydrolysis was observed as a significant mechanism of degradation at higher temperatures [35]. In another study, addition of Tween 80® inhibited aggregation of IL-2 mutein during shaking. Paradoxically, Tween 80® accelerated the aggregation of the same protein in a temperature-dependent manner during storage. The build-up of peroxides from autoxidation of degraded polysorbates increased the oxidization rate of the protein, therefore compromising its stability in storage [36]. 2.2.2.2. Poloxamers Triblock copolymers of the form polyethylene oxide–polypropylene oxide–polyethylene oxide (PEO–PPO–PEO), or poloxamers (commercially available as Pluronics® or SynperonicsTM), constitute another class of nonionic surfactant commonly used in the pharmaceutical industry (Figure 2.3). The poloxamers are listed as pharmaceutical excipients in the U.S. and British Pharmacopoeia, and have been used extensively in a variety of pharmaceutical formulations [37]. Poloxamers show complex aggregation behavior, involving unimers, oligomers, micelles of various geometries, and larger clusters, with strong temperature and concentration dependences. The critical micelle temperature and CMC values of such triblocks have been estimated over a wide 20 range of molecular weights and PPO/PEO ratios, by a number of different methods [26, 38-41]. In general, triblocks with a larger hydrophobic (PPO) domain form micelles at lower concentrations or, at a constant triblock molar concentration, have lower critical micelle temperatures. For a given PPO:PEO ratio, triblocks of higher molecular weight form micelles at lower concentrations and temperatures. The size of the hydrophilic PEO group appears to play a smaller role in the micellization process. Alexandridis et al. [39Error! Bookmark not defined.] performed a thermodynamic analysis of the formation of triblock micelles, to obtain standard free energies, enthalpies, and entropies of micellization for a number of poloxamers. They found that the standard enthalpy of micellization was positive for all triblocks tested, indicating that the transfer of unimers from solution to the micelle is an enthalpically unfavorable, endothermic process. A negative entropy contribution was thus implicated as the driving force for micellization. Poloxamer 188 (BASF Pluronic® F68, Figure 2.3) is widely used for the large scale production of mammalian cell culture, and also where bioreactors are increasingly used to amplify a cell population. It is used as a shear-protective excipient to enhance cell yield in agitated cultures and reduce cell adhesion in stationary cultures [42]. Two mechanisms have been proposed in the literature to explain the cell protection effect of poloxamer 188. One suggests that it affects the culture medium characteristics, by inhibiting damage associated with cell-bubble interactions when, for example, a bubble breaks at the air-water interface. Another suggests that cells exhibit higher resistance to shear stress in the presence of the triblocks. Poloxamer 188 has also been reported to facilitate the refolding and to suppress aggregation of a thermally denatured protein [43]. However, removing poloxamer 188 during product recovery may compromise the product yield, as well as inhibit the growth of some cell lines [44]. 21 HO O H O O 79 CH3 26 79 Figure 2.3: Chemical structure of the PEO-PPO-PEO triblock copolymer Poloxamer 188. Similar products with various molecular weights and PEO: PPO ratios are also commercially available [40]. 2.2.3. Surfactant modulation of protein adsorption and aggregation Surfactants stabilize proteins by two major mechanisms: (a) by preferentially locating at an interface, in this way precluding protein adsorption, and/or (b) by associating with proteins in solution, in this way stabilizing them against close approach and inhibiting aggregation (Figure 2.4). Some surfactants may function according to only one of these mechanisms, while others may function according to both. protein protein protein a b Figure 2.4: Mechanisms of stabilization of proteins by surfactants, which may (a) dominate the interface and prevent protein adsorption, or (b) preferentially associate with proteins and thus prevent close approach and aggregation. 2.2.3.1. Protein stabilization by surfactant adsorption at interfaces A number of experimental investigations of the interfacial behavior of surfactant and protein mixtures have been conducted, and these have identified three possible adsorption outcomes: complete hindrance, reduced amounts, or increased amounts of protein adsorption. Complete hindrance is attributed to the faster diffusion of the (generally smaller) surfactant molecules to the interface, as compared to the much 22 larger protein molecules. The adsorbed surfactant layer coats the interface, and sterically prevents protein adsorption. Reduced or increased amounts of adsorption are usually attributed to the formation of surfactant-protein complexes with reduced or increased surface affinity, respectively. In either case, the behavior of these complexes is considerably different from that of the pure protein or surfactant in solution. An important goal in biotechnology process development and biopharmaceutical formulation engineering is to minimize the protein loss that occurs throughout the process by colloidal and interfacial mechanisms, e.g., aggregation and adsorption [45, 46]. In order to achieve this, a fundamental understanding of the mechanisms underlying surfactant effectiveness is necessary. In particular, better understanding of the specific roles of the surfactant, protein, and surfactant-protein complex in modulating interfacial behavior will provide direction for much-needed process improvements in the production and finishing of therapeutic proteins. 2.2.3.1.1. Expectations based on sequential and competitive adsorption experiments The sequential introduction of a surfactant following protein adsorption at an interface may result in the removal of adsorbed protein, due to the formation of surfactant-protein complexes and subsequent solubilization of these complexes. Alternatively, adsorbed protein may be displaced by surfactant on account of a stronger surfactant-surface association. The extent of surfactant-mediated removal of adsorbed protein depends on protein, surfactant and surface properties, and also other factors [47]. In general, the difference in the amount of adsorbed protein eluted by anionic, cationic and nonionic surfactants correlates with the strength of binding between the surfactant and the protein in solution [48]. Nonionic surfactants, which are known to bind rather weakly to proteins, are least effective in removing adsorbed protein molecules from the 23 interface. In particular, when introduced to an adsorbed protein layer on a hydrophilic surface, nonionic surfactants generally have little effect on the adsorbed amount. In contrast, on hydrophobic surfaces, nonionic surfactants typically have a substantial effect on the adsorbed protein, presumably because of the difference in surfactant binding strength at the interface [49]. Joshi and McGuire [50] have described the interaction of lysozyme, a well-characterized globular protein, with the nonionic surfactant polysorbate 80 at solidwater interfaces. The concentration of the surfactant, as well as the method of surfactant and protein introduction to the surface (i.e. sequential or combined) was varied in order to elucidate the separate roles of protein, surfactant, and the protein-surfactant complex in determining adsorption outcomes. They reported a decrease in lysozyme adsorption on hydrophobic silica upon addition of polysorbate 80, and this reduction in adsorbed protein increased with the concentration of polysorbate 80 in solution. Sequential adsorption experiments showed that, at sufficiently high concentration, polysorbate 80 was able to remove adsorbed lysozyme from a hydrophobic surface. In addition, if polysorbate 80 was introduced to the hydrophobic surface prior to addition of lysozyme, adsorption of the protein was reduced or even eliminated. On the other hand, polysorbate 80 had no effect on the adsorption of lysozyme onto hydrophilic silica. Finally, sequential adsorption experiments showed that polysorbate 80, when introduced to the interface either before or after adsorption of lysozyme, had no effect on the amount of lysozyme adsorbed. The observed differences in protein adsorption were attributed to surfacedependent differences in the binding affinity of polysorbate 80 to hydrophobic or hydrophilic surfaces; this work emphasizes the importance of direct interactions between the surfactant and the solid surface, relative to surfactant-protein interactions in solution. Accordingly, the rapid diffusion of the small surfactant molecules to the interface 24 (relative to proteins) is likely to contribute to a reduction in protein adsorption only if the surfactant-surface affinity is sufficiently high. 2.2.3.1.2. On the stabilization of rFVIII by polysorbate 80 at solid-water interfaces Lysozyme is a much-used “model” protein for the study of adsorption phenomena in a number of well-controlled circumstances, but results of such work have contributed to forming a foundation for the greater understanding of the behavior of more complex therapeutic proteins in surfactant-containing formulations. The adsorption, structural alteration and biological activity of a recombinant Factor VIII (rFVIII) was investigated at a hydrophilic and hydrophobic solid-water interface in the presence of polysorbate 80 [24]. At the hydrophobic surface, the presence of polysorbate 80 in the protein solution resulted in reduced protein adsorption, while rFVIII adsorption at the hydrophilic surface was entirely unaffected by the presence of polysorbate 80. As in the case of polysorbate 80 and lysozyme, these observations were attributed to the high binding strength between the surfactant and the hydrophobic surface, and the relatively low affinity between polysorbate 80 and the hydrophilic surface. Association of polysorbate 80 and rFVIII in solution was observed to be entirely ineffective in reducing rFVIII adsorption, indicating that the surfactant prevents adsorption by coating the interface, not the individual protein molecules. In the absence of surfactant, proteins can be expected to adsorb with high affinity to hydrophobic surfaces as well as negatively-charged, positively-charged, and electronically neutral surfaces [51]. Substantial reductions in protein adsorption can be observed with surfactant addition under appropriate circumstances, or in general through the application of so-called “nonfouling” coatings, such as those exhibiting pendant PEO chains [52-55]. The pendant polymer chains resist protein adsorption by several 25 mechanisms, primarily steric repulsion [56]. It is thus reasonable that steric repulsion is a requirement for eliminating protein adsorption, and explains the nonfouling effect of such coatings. In the context of the work summarized in relation to polysorbate 80, steric repulsion adequately explains the protein-repellent effect of added surfactants in the presence of surfaces for which surfactant-surface binding is strong, and the absence of any significant effect of added surfactant with surfaces for which the surfactant-surface binding is weak. It follows that steric repulsion is a requirement for the surfactantmediated prevention of protein aggregation. 2.2.4. Protein stabilization by surfactant-protein association Protein stabilization by association of surfactants requires only sufficiently strong surfactant-protein interaction, and would be effective in reducing protein adsorption, regardless of the strength of surfactant-surface binding. It is instructive to consider this mechanism of protein stabilization, with reference to results of surfactant action recorded at air-water interfaces in the presence and absence of proteins. 2.2.4.1. A simple view of surfactant-protein mixtures at the air-water interface The molecular dynamics contributing to changes in air-water interfacial tension for protein-surfactant mixtures are complex, but offer an insight into the mechanisms of surfactant interactions with interfaces and proteins. In ideal circumstances (i.e., for random chain protein molecules and small, ionic surfactants) the following equilibrium behavior is expected with increasing surfactant concentration (Figure 2.5) [57, 58]: 26 Region 1: At very low surfactant concentrations, the steady-state interfacial tension is the same as it would be for a pure protein solution. The relatively few surfactant molecules have little or no effect on surface tension. Region 2: As surfactant concentration increases, the interfacial tension decreases, due to surfactant occupation of “empty sites” at the air-water interface, as well as the formation of surface-active protein-surfactant complexes. Region 3: At higher surfactant concentrations, the interfacial tension is expected to plateau, presumably because it is energetically favorable for surfactant to bind to protein at these concentrations (in this range, the CMC recorded for the surfactant in protein-free buffer may be exceeded). Region 4: Equilibrium interfacial tension decreases again with increasing surfactant concentration, as a result of complete displacement of protein from the interface by the surfactant. Region 5: Further increases in surfactant concentration have no effect on interfacial tension, and a second plateau is reached. In this regime, the CMC, which is specific to the protein concentration used in the experiment, has been reached, and no further surfactants can adsorb to the air-water interface [59]. 27 2 3 4 5 Steady-State Surface Tension 1 Region 1: Very low surfactant concentrations air liquid protein • Surfactant predominantly appears as unimers in the bulk liquid phase • Limited interactions of surfactant molecules with protein and liquid interface Surfactant Concentration (log10) Region 2: Low surfactant concentrations air liquid protein • Surfactant unimers appear at interface and in bulk liquid Region 5: Very high surfactant concentrations (> CMC) air • Surfactant molecules begin to displace protein from the air-liquid interface protein • Appreciable interactions of surfactant molecules with liquid interface and protein liquid • Substantial interactions of surfactant molecules with liquid interface and protein air liquid • Surfactant unimers appear at interface, in bulk liquid, and at protein surfaces • Appreciable interactions of surfactant molecules with liquid interface, but limited interaction with protein Region 4: High surfactant concentrations Region 3: Moderate surfactant concentrations protein • Protein completely displaced from air-liquid interface • Strong interactions between surfactant and interface; protein coated by surfactant protein • Micelles form in bulk liquid Figure 2.5: Schematic illustration of surface tension depression associated with five regimes of protein-surfactant and surfactant-interface interactions. Redrawn with permission from [58]. This description relates to an idealized protein-surfactant mixture at equilibrium, and is a useful reference for interpreting observations of systems of greater complexity. However, for theoretical and practical reasons, measurements of the true interfacial equilibrium are usually not possible for real protein-surfactant mixtures. This is due mainly to the inherent irreversibility of protein adsorption, as well as uncertainties in the measurements required by the experiments. The interfacial tension kinetic and steady-state behaviors exhibited by surfactant solutions, as a function of surfactant concentration, and in the presence and absence of protein, have been recorded for a number of systems. Comparison of the steady-state surface tension recorded for surfactant-protein mixtures with that of surfactant alone (at similar concentrations) can be used to reveal whether protein 28 adsorption is evident, or if the steady-state interfacial behavior is governed entirely by surfactant. In particular, if no appreciable difference is recorded between the steady-state value of interfacial tension demonstrated by protein-surfactant mixtures and by the surfactant alone at a similar concentration, one may tentatively conclude that only the surfactant undergoes appreciable adsorption at the interface (Figure 2.6). For example, air-water interfacial tensiometry experiments were performed with mixtures of rFVIII and polysorbate 80. The measured steady-state interfacial tensions, with and without the protein, were identical at surfactant concentrations above 18 ppm, indicating that the Surface Tension (γ) surface was dominated by the surfactant [22]. Low [S] S only S+P Intermediate [S] High [S] S only S+P S/S+P Time Figure 2.6: Schematic of differences in kinetics and extent of surface tension depression by surfactant alone (S) or surfactant with protein (S + P), at different surfactant concentrations ([S]) for a constant protein concentration. In general, surface tension depression increases faster at a given [S] in the presence of protein. Steady-state surface tensions corresponding to S and S + P converge at sufficiently high [S]. 2.3. A testable, thermodynamic argument to guide surfactant selection 2.3.1. Insights gained from intact and protein-depleted pulmonary surfactant Schram and Hall [60] determined the influence of the two hydrophobic proteins, SP-B and SP-C, on the thermodynamic barriers that limit the adsorption of pulmonary surfactant vesicles to the air–water interface in the lung. Vesicle adsorption, in 29 this case, is characterized by separation of the surfactant acyl chains, followed by fusion of the bilayer vesicle with the interface to form a monolayer (Figure 2.7). For this purpose they measured the kinetics of adsorption (based on interfacial tensiometry) for intact calf lung surfactant extract, and compared them with adsorption of an extract containing the complete set of surfactant lipids, but depleted of the SP-B and SP-C proteins. The surfactant proteins SP-B and SP-C are critical for normal respiration, and accelerate the adsorption of intact surfactant (relative to proteinfree surfactant) more than ten-fold. This physiological behavior was accurately reflected in the surface tension kinetic results recorded by Schram and Hall. They interpreted their kinetic results for intact and protein-free surfactant adsorption with reference to a mechanism for vesicle adsorption. They postulated that vesicle adsorption and fusion is governed by the formation of a rate-limiting structural intermediate between the free and adsorbed forms (Figure 2.7). Air Water Figure 2.7: Schematic of the rate-limiting intermediate state in the transition from a phospholipid bilayer to an interfacial monolayer. Proteins located within the bilayer were found to reduce the enthalpic barrier of the intermediate state. Adapted from [60] with permission from Elsevier. 30 In particular they measured the rate constant (km) characterizing the slope of the surface tension–time isotherm during the initial decrease in surface tension, at each of a series of different temperatures and concentrations, according to: rate = km ⋅ c n , (1) where n, the order of the reaction, was obtained from measurements of the initial adsorption rate at each concentration, c, in the bulk phase. The activation energies, Ea, for adsorption were then derived from the slopes of plots of the experimental ln km vs. 1 T data, according to the Arrhenius equation: E 1 ln km = − a ⋅ + ln A , R T (2) where R is the gas constant, T is temperature, and A is the Arrhenius pre-exponential factor. They invoked transition-state theory to consider the expected effect of temperature, in terms of an equilibrium between the “reactants” (i.e., the vesicles and unoccupied airwater interface) and an activated complex. From this model, the rate constant can be described in thermodynamic terms: k T km = b h −∆G RT , e (3) where kb and h are Boltzmann’s and Planck’s constants, respectively, and ΔG is the Gibbs free energy of transition. Thus, since ∆G = ∆Η − Τ∆S , k k ∆S ∆H 1 − ⋅ + ln b + ln m = R R T T h (4) 31 The slope and intercept of plots of ln km T vs. 1 T therefore provide quantitative estimates of the enthalpy (ΔH) and entropy (ΔS) of the transition. The transition of a bilayer to form an interfacial monolayer requires a transient exposure of the hydrophobic tails of the surfactant lipids to the aqueous environment (Figure 2.7), and consequently has an unfavorable entropy of transition. Schram and Hall’s analysis, however, showed that the surfactant proteins did not affect the entropy of transition; rather, the essential effect of the proteins was to minimize an unfavorable enthalpy barrier to formation of the structural intermediate. This enthalpic cost was attributed to the dissociation of the surfactant acyl chains during the separation of the leaves of the bilayers. An interesting observation characteristic of surface tension depression by surfactant-protein mixtures is that the kinetics of surface tension depression in such mixtures tend to be uniformly greater than that recorded for surfactant alone at the same concentration, regardless of whether the final steady-state surface tension is similar in each case (Figure 2.6). This kind of “synergistic” effect is well documented for synthetic polymer-surfactant mixtures [61], but less well understood in relation to proteinsurfactant mixtures. Joshi et al. [22] found the rate of surface tension decrease to be greater for polysorbate 80-rFVIII mixtures than for polysorbate acting alone, at all polysorbate concentrations studied in that work (8 to 108 ppm). The reasons for this have not been articulated in any quantitative fashion, but might be explained in part using an approach to the problem similar to that outlined by Schram and Hall [60]. This approach might also provide direction for surfactant selection (or surfactant design) to more effectively manage issues surrounding aggregation and adsorption loss. 32 2.3.2. Surfactant-protein association at surfactant concentrations above the CMC Consider the case of surfactant adsorption in the presence of protein, at surfactant concentrations above the CMC (or otherwise consistent with the presence of associated unimers, if the surfactant system is not governed by any obvious CMC). The hydrophobic interactions that maintain the structure of a micelle or other surfactant aggregate are based on entropy, resulting from the highly-ordered clathrate cages of water molecules that surround a nonpolar molecule in an aqueous environment. A thermodynamic analysis similar to that used by Schram and Hall [60] can be applied to the notion of protein-mediated acceleration of surfactant adsorption introduced in Section 3.1. We speculate that proteins may accelerate the adsorption of surfactants by reducing the major entropic barrier faced by the surfactant in moving from the aggregate to interface, and/or by reducing the enthalpy of activation. In the latter case, the proteins might destabilize the surfactant self-association, or they could produce a “catalytic” reduction in the enthalpy of some structural intermediate between unbound surfactant aggregates and adsorbed surfactant unimers (Figure 2.8). In either case, the likely source for a change in enthalpy would be an alteration of the van der Waals interactions among the regions on the surfactant molecules mediating their self-association. Separation of surfactant monomers from aggregates and their location at the air-water interface would disrupt van der Waals interactions and produce an unfavorable enthalpy. 33 air water + air water + + protein protein protein Figure 2.8: Hypothetical mechanisms for transport of surfactant from an aggregated state to the surface. In the absence of protein (top), surfactant unimers must dissociate and migrate through the liquid to the interface. Proteins may promote formation of a structural intermediate between surfactant aggregates and adsorbed unimers, reducing the thermodynamic barriers associated with their location at the interface (bottom). These arguments may be more significant in relation to associations of large-molecule surfactants (e.g. poloxamers), as opposed to the smaller surfactants. The mechanism of adsorption and the adsorption kinetics exhibited by poloxamers, for example, strongly depends on their solution concentration during adsorption [62, 63]. At low concentrations, the triblocks exist as individual molecules (unimers), and their adsorption may not uniformly cover the entire available surface. At high concentrations, however, adsorption is dominated by micelles or other aggregates, and the PEO chains may inhibit the association between the hydrophobic center blocks and the surface. Adsorption of poloxamers is slow in comparison to small molecule surfactants, yet a synergistic effect in the kinetics of surface tension depression similar to that described in Section 2.4.1 has been recorded for mixtures of selected proteins and poloxamers 188 (Pluronic® F68). As with the polysorbates, surface tension depression in a poloxamerprotein mixture tends to be faster than for the same concentration of the surfactant alone (Figure 2.6). 34 2.3.3 Surfactant-protein association at surfactant concentrations below the CMC There is convincing evidence that poloxamers interact with the plasma membranes of cells. More hydrophobic poloxamers generally show greater tendencies to incorporate into the cell plasma membrane [63]. Gigout et al. [64] examined the incorporation of poloxamer 188 into the cell plasma membrane and its subsequent uptake by chondrocytes and CHO cells. They were able to conclusively demonstrate that the triblocks did in fact enter the cells, and possibly accumulate in the endocytic pathway. While poloxamer 188 shows a high affinity for entropically-driven associations with cell membranes, it is not expected to form micelles in water except at very high concentrations [38, 40, 41, 65]. It is possible that the observed high affinity for cell surfaces can be explained by the significantly decreased solubility of the surfactant in the presence of salts. Patel et al. studied the micellization of a very hydrophilic poloxamer (80% PEO) in various sodium salt solutions. While the triblock did not form micelles in pure water at ambient temperatures, it exhibited aggregation and micellization in salt solutions. The cloud point and critical micelle temperature (CMT) decreased substantially in the presence of salts. They found that the average micelle size increased with salt concentration, and the relative effect of the different anions to enhance micellization followed the Hofmeister series (i.e. PO43– > SO42– > F– > Cl– > Br–), consistent with salting out of the poloxamer [66]. Clouding is a thermodynamic phase transition that is characteristic of molecules containing PEO or other polyethers, and is commonly associated with triblocks and other nonionic surfactants in water. It is due in part to changes in hydrogen bonding and increasingly attractive interactions between the PEO chains as the temperature increases, making the water a “less good” solvent [67]. The influence of salts can be 35 explained in a similar fashion, being primarily caused by the existence of a salt-deficient zone surrounding the PEO chains: small ions with little polarizability would be repelled by the poorly polarizable PEO chain. This salt-deficient zone gives rise to an attractive component in the interaction between PEO segments. As polymer segments approach each other, their salt depletion zones overlap and the surrounding water molecules are liberated to the bulk solution. The excluded water molecules have a lower chemical potential in the bulk, a thermodynamically favored state [38]. Cell surfaces (as well as the “surfaces” of proteins) are charged and “separated” from the bulk solution by a surrounding ion-enriched electrical double layer [68,69]. It is reasonable to anticipate a marked decrease in solubility of a poloxamer at the vicinity of such a colloidal surface, owing to the higher salt concentration at the interface than in the bulk. It is thus tempting to hypothesize that a surfactant unimer that approaches a cell or protein “surface” would find itself in a region of higher ionic strength, and be driven to associate with the surface by a “salting out” mechanism. Thus, surfactants could effectively stabilize a protein by association with the surface, forming a surfactant-protein complex that guards against close approach and subsequent aggregation with other proteins. Conversely, the surface at an air-water interface is depleted of salts relative to the bulk solution [70], and surfactant unimers at the surface would encounter a “more watery” micro-environment, and would thus not adsorb with high affinity to the interface. The effects of these microscopic changes in ionic strength are consistent with the accelerated surface tension depression observed for surfactantprotein mixtures (relative to protein-free surfactant), and also with protein stabilization in the vicinity of the air-water interface, even when the interface is only partially occupied by surfactant. Further research into these mechanisms is clearly warranted. 36 2.4. Conclusions Surfactants stabilize proteins by their preferential location at an interface and/or their association with protein in solution. In the common case of surfactant-protein interaction governed by hydrophobic association, the molecular origins of surfactantmediated stabilization of protein can be inferred, in part, from quantitative models. Insights can be gained by comparison of thermodynamic barriers that limit surfactant adsorption, from surfactant-protein mixtures and protein-free surfactant solutions, to the air–water interface. These barriers are defined by the enthalpy and entropy of transition from free (whether unimeric or aggregated) surfactant molecules to adsorbed surfactants, via formation of a rate-limiting structural intermediate. This is easily done, for example, through analysis of surface tension kinetic data recorded at different temperatures. 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Lett. 103 (2009) 257802-257805. 42 PROTEIN EFFECTS ON SURFACTANT ADSORPTION SUGGEST THE DOMINANT MODE OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN Hyo Jin Lee,a,b Arnold McAuley,b and Joseph McGuirea,* a School of Chemical, Biological and Environmental Engineering, Oregon State University, Corvallis, OR 97331 b Proces and Product Development Department Amgen Inc., Thousand Oaks, CA 91320 * Corresponding Author: Joseph McGuire School of Chemical, Biological and Environmental Engineering Oregon State University 103 Gleeson Hall Corvallis, OR 97331-2702 Tel: 541-737-6306 Fax: 541-737-4600 Email: mcguirej@engr.orst.edu 43 CHAPTER 3 PROTEIN EFFECTS ON SURFACTANT ADSORPTION SUGGEST THE DOMINANT MODE OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN Abstract Nonionic surfactants are commonly used to stabilize proteins in upstream and downstream processes and drug formulation. However, the effectiveness of a surfactant strongly depends on its mechanism(s) of action but selection of surfactants for protein stabilization currently is not made with benefit of any quantitative, predictive information to ensure that this requirement is met. Often the kinetics of surface tension depression (by surfactant) in protein-surfactant mixtures tends to be greater than that recorded for surfactant alone at the same concentration. Here, we compare surface tension depression by poloxamer 188 (Pluronic® F68), polysorbate 80 (PS80), and polysorbate 20 (PS20) in the presence and absence of lysozyme and recombinant protein, at different surfactant concentrations and temperatures. The kinetic results are interpreted with reference to a mechanism for surfactant adsorption governed by the formation of a rate-limiting structural intermediate (i.e., an “activated complex”) between surfactant aggregates and protein. The presence of lysozyme was seen to increase the rate of surfactant adsorption in relation to surfactant acting alone at the same concentrations for the polysorbates while less of an effect was seen for Pluronic® F68. However, the addition of salt was observed to accelerate the surface tension depression of Pluronic® F68 in the presence of lysozyme. Findings of aggregate present in polysorbates even below the CMC and Pluronic® F68 displaying greater aggregation and micellization in salt solutions have lead us to believe that an interaction between lysozyme and surfactant 44 aggregates may cause aggregate destabilization and an increased availability of surfactant monomers for adsorption. The addition of a more hydrophobic, surface active protein in place of lysozyme resulted in greater enhancement of surfactant adsorption than that recorded in the presence of lysozyme. A simple thermodynamic analysis indicated the presence of protein caused a reduction in ∆𝑮 for the surfactant adsorption process, with this reduction deriving entirely from a reduction in ∆𝑯. We suggest that protein accelerates the adsorption of these surfactants by disrupting their self associations, releasing surfactant monomers. 3.1 Introduction Aggregation phenomena in protein therapeutics have been studied and reported extensively by academia, industry and regulatory agencies. Aggregation is highly undesirable due to the profound impact on the stability of the drug product, which can result in loss of activity, unwanted immunogenic responses, and increased rate of rejection as a marketable product [1]. Several workers have reported on the different mechanisms of aggregation, and suggested possible methods to inhibit aggregation [2-4]. Several mitigation strategies are used in the biotechnology industries to stabilize therapeutic proteins, but the addition of surfactants appears to be a general approach [5]. Surfactants are able to stabilize proteins through two major mechanisms. One involves the preferential location of surfactant at nearby interfaces, in this way precluding protein adsorption [6]. The other mechanism involves association of protein and surfactant molecules into complexes that prevent proteins from interacting with surfaces as well as each other [7]. In general, both mechanisms must be at play for effective protein stabilization against aggregation and activity loss, but selection of 45 surfactants for protein stabilization currently is not made with benefit of any quantitative, predictive information to ensure that this requirement is met. An interesting observation characteristic of surface tension depression by surfactant-protein mixtures is that the kinetics of surface tension depression (by surfactant) in such mixtures often tends to be greater than that recorded for surfactant alone at the same concentration [8]. It is reasonable to expect that such “accelerated adsorption” of surfactant in the presence of protein is governed by surfactant-protein interactions and must therefore be a function of parameters important to protein stabilization by that surfactant. We recently hypothesized that parameters determining whether surfactant stabilization occurs mainly by association with the interface or by association with protein ought to be quantifiable for different surfactants by analysis of such accelerated adsorption data [9]. In particular, the adsorption of surfactant monomers may be enhanced by interaction between surfactant aggregates and protein near the interface, leading to the release of monomers for adsorption. As depicted in Figure 3.1 (top), transport of monomers from an aggregated state to the air-water interface in the absence of protein would require that they dissociate from the aggregate and migrate through the liquid in order to adsorb. The presence of protein may reduce the major enthalpic barrier associated with surfactant dissociation, and/or reduce the major entropic barrier faced by the surfactant in moving from the aggregate to the interface [10]. The protein may facilitate aggregate disruption, leading either to an increased concentration of surfactant monomers thus enhancing adsorption (Figure 3.1, middle), or to the formation of stable, surfactant-protein complexes having little or no effect on surfactant adsorption rate (Figure 3.1, bottom). Thus we expect that accelerated surfactant adsorption in the presence of protein will occur with surfactants that stabilize protein mainly by their own 46 adsorption at interfaces, and we expect the absence of accelerated surfactant adsorption will occur with surfactants that form stable surfactant-protein associations. air water + air water + protein protein protein air water protein protein protein Figure 3.1: In the absence of protein (top), surfactant monomers must dissociate and migrate through the liquid in order to adsorb. The presence of protein may facilitate aggregate disruption, leading either to an increased concentration of surfactant monomers thus enhancing adsorption (middle), or to the formation of stable, surfactant-protein complexes having little or no effect on surfactant adsorption rate (bottom). In this paper we complete the first step in testing the hypothesis outlined above, through comparison of surface tension depression by selected surfactants in the presence and absence of protein, at different surfactant concentrations and temperatures. The kinetic results are interpreted with reference to a mechanism for surfactant adsorption governed by the formation of a rate-limiting structural intermediate (i.e., an “activated complex”) between surfactant aggregates and protein. 47 3.2 Materials and Methods 3.2.1. Proteins and surfactants The proteins used were lyophilized chicken egg white lysozyme (Sigma Aldrich, Saint Louis, MO) and an Amgen recombinant protein. Lysozyme was dissolved in Dulbecco’s phosphate buffered saline (1X PBS, Invitrogen, Carlsbad, CA) at 100 mg/mL or higher and was filtered using a 0.2 um PVDF filter (Millipore, Billerica, MA) before the surface tension experiments. The final concentration of lysozyme used in the measurements was 5 mg/mL. Amgen recombinant protein is 18.8 kDa that consists of 174 amino acids with 2 disulfide bonds. Recombinant protein was provided in 10 mM sodium acetate, 5% sorbitol at pH 3.5 at 3 mg/mL and was concentrated to 20 mg/mL using Amicon stir cells (Millipore, Billerica, MA) and filtered using a 0.2um PVDF filter before use. The final concentration of the recombinant protein used in the measurements was 1mg/mL. The surfactants studied were Poloxamer 188 (Pluronic ®F68, BASF, obtained from Sigma Aldrich, Saint Louis, MO) polysorbate 80 (PS 80, Croda, East Yorkshire, England) and polysorbate 20 (PS 20, Croda, East Yorkshire, England). Stock solutions at high concentrations of the surfactants were made in 1X PBS and in 10 mM sodium acetate, 5% sorbitol at pH 3.5 buffers and diluted to the respective buffers to have a range of concentrations. Buffer salts and sorbitol were from obtained from JT Baker (Philipsburg, NJ) 3.2.2. Surface tension measurements Surface tension was measured using a 1-cm wide Wilhelmy plate (Kruss, Hamburg, Germany) on a Kruss K100 tensiometer (Hamburg , Germany). 19 mL of buffer was placed in a 50 mL glass vessel housed within a temperature controlled jacketed stage and surface tension was recorded every 10 seconds for 220 seconds. Then, 48 1 mL of the desired sample (which could be the protein or surfactant by itself or surfactant-protein mixture) was introduced and stirred for 4 seconds before measurement started again and recorded every 10 seconds for a total of 220 seconds and repeated 3 times. In order to capture the initial surface tension depression kinetics, only the last data point of the buffer is shown at time zero. Most of the experiments were done at room temperature. However to study the effect of temperature on surface tension depression, additional experiments were performed at 5 °C and 40 °C. 3.3 Results and Discussion 3.3.1. Effect of surfactant concentration Representative plots of the early kinetics of surface tension depression are shown in Figures 3.2-3.4 for selected concentrations of PS 80 (Figure 3.2), PS 20 (Figure 3.3), and Pluronic ® F68 (Figure 3.4), in the presence and absence of lysozyme. Both the initial rate and extent of surface tension depression generally increased with increasing surfactant concentration. Lysozyme alone at 5 mg/mL displayed less surface tension depression (about 64 mN/m after 220 s) than any of the surfactants at surfactant concentrations of 5 ppm and above. Thus the plots of Figures 3.2-3.4 suggest that for lysozyme-surfactant mixtures, interfacial behavior was governed predominantly by surfactant at concentrations greater than 5 ppm. 49 75 Surface Tension (mN/m) 70 5 ppm PS 80 5 ppm PS 80 +Lysozyme 10ppm PS 80 10 ppm PS 80 +Lysozyme 20 ppm PS 80 20 ppm PS 80 +Lysozyme 50 ppm PS 80 50 ppm PS 80 +Lysozyme 100 ppm PS 80 100 ppm PS 80 +Lysozyme 5 mg/mL Lysozyme 65 60 55 50 45 40 0 50 100 150 200 250 Time (sec) Figure 3.2: Adsorption isotherm (surface tension vs. time) of PS 80 and lysozyme mixture at 25 °C in 1X PBS. 75 Surface Tension (mN/m) 70 5 ppm PS 20 5 ppm PS 20 +Lysozyme 10 ppm PS 20 10 ppm PS 20 +Lysozyme 20 ppm PS 20 20 ppm PS 20 +Lysozyme 50 ppm PS 20 50 ppm PS 20 +Lysozyme 100 ppm PS 20 100 ppm PS 20 +Lysozyme 5 mg/mL Lysozyme 65 60 55 50 45 40 35 0 50 100 150 200 250 Time (sec) Figure 3.3: Adsorption isotherm (surface tension vs. time) of PS 20 and lysozyme mixture at 25 °C in 1X PBS. 50 75 Surface Tension (mN/m) 70 5 ppm F68 5 ppm F68 +Lysozyme 10 ppm F68 10 ppm F68 +Lysozyme 20 ppm F68 20 ppm F68 +Lysozyme 50 ppm F68 50 ppm F68 +Lysozyme 500 ppm F68 500 ppm F68 +Lysozyme 5 mg/mL Lysozyme 65 60 55 50 45 0 50 100 150 200 250 Time (sec) Figure 3.4: Adsorption isotherm (surface tension vs. time) of Pluronic® F68 and lysozyme mixture at 25 °C in 1X PBS. For PS 80 and PS 20 at the lowest concentrations shown (5 and 10 ppm), the presence of lysozyme was seen to increase the rate of surfactant adsorption in relation to surfactant acting alone at the same concentrations. Using the dye 1,6-diphenyl 1,3,5hexatriene, Kerwin [13] recorded initiation of aggregate formation in solutions of polysorbates 20 and 80 at concentrations well below the CMC of each surfactant, with PS 80 showing more pronounced aggregation than PS 20. In such tests, an increase in fluorescence is detected when the (hydrophobic) dye partitions from the aqueous phase into the apolar phase of an aggregate interior [11,12]. Both polysorbates showed initiation of aggregate formation in that work at concentrations consistent with those used in construction of Figures 3.2-3.3. The enhanced surfactant adsorption recorded in the presence of lysozyme (Figures 3.2 and 3.3) is thus qualitatively consistent with an interaction between lysozyme and surfactant aggregates causing aggregate destabilization and an increased availability of surfactant monomers for adsorption. It is also consistent with the observation that accelerated surfactant adsorption is more evident for PS 80 than 51 for PS 20 in these cases. At polysorbate concentrations of 20 ppm and above, no appreciable difference was recorded between lysozyme-surfactant mixtures and the surfactant alone at the same surfactant concentration. This suggests that the population of surfactant monomers near the interface is sufficiently high (and the strength of proteinaggregate association sufficiently weak), such that adsorption is no longer rate-limited by the protein-aggregate structural intermediate. On the other hand, lysozyme showed much less of an effect on Pluronic® F68 adsorption. This would be consistent with Pluronic® F68 undergoing no appreciable aggregate formation under these conditions. Pluronic® F68 is not expected to form micelles in water except at very high concentrations [14-17]. Like other Pluronics® it shows decreased solubility in the presence of salts. Thus we would expect lysozyme to show a more pronounced effect on Pluronic® F68 adsorption in the presence of salt. 3.3.2. Effect of salt While Pluronic® F68 would exhibit greater aggregation and micellization in salt solutions, the surface at an air-water interface is depleted of salts relative to the bulk solution [18], and surfactant monomers would thus not be expected to adsorb with high affinity to the interface. Figure 3.5 shows a negligible effect on adsorption of Pluronic® F68 in the absence of lysozyme, upon addition of 250 mM NaCl. However, the addition of salt did accelerate the surface tension depression of Pluronic® F68 in the presence of lysozyme, albeit modestly, consistent with an increased population of Pluronic® F68 aggregates in solution. Even at 50 ppm, where Figures 3.2-3.4 indicate the adsorption is apparently not rate-limited by the protein-aggregate structural intermediate for any of the three surfactants studied, the addition of salt was observed to accelerate the surface tension depression of Pluronic® F68 in the presence of lysozyme 52 (Figure 3.6). No effect upon addition of salt was evident for adsorption of PS 80 or PS 20. This is consistent with prior reports of polysorbate aggregation behavior being largely unaffected by the presence of salts [13,19]. 76 74 Surface tension (mN/m) 72 70 68 10 ppm F68 0 mM NaCl 10 ppm F68 +Lysozyme 0 mM NaCl 10 ppm F68 250 mM NaCl 10 ppm F68 +Lysozyme 250 mM NaCl 66 64 62 60 58 56 54 0 50 100 150 200 250 Time (sec) Figure 3.5: Effect of salt on the adsorption isotherm (surface tension vs. time) of 10 ppm Pluronic® F68 and lysozyme mixture 75 Surface tension (mN/m) 70 65 50 ppm F68 250 mM NaCl 50 ppm F68 +Lysozyme 250 mM NaCl 50 ppm PS 80 250 mM NaCl 50 ppm PS 80 + Lysozyme 250 mM NaCl 50 ppm PS 20 250 mM NaCl 50 ppm PS 20 + Lysozyme 250 mM NaCl 60 55 50 45 40 0 50 100 150 200 250 Time (sec) Figure 3.6: Effect of salt on the adsorption isotherm (surface tension vs. time) of surfactants (50 ppm Pluronic® F68, PS 80 and PS 20) and lysozyme mixture 3.3.3. Effect of temperature and comparison to a simple model for protein-mediated acceleration of surfactant adsorption 53 The early kinetics of surface tension depression were recorded for each surfactant at 5 and 10 ppm, in the presence and absence of lysozyme, at 5, 25, and 40 °C. Representative plots (constructed for surfactant concentrations of 5 ppm) are shown in Figures 3.7-3.9 for PS 80 (Figure 3.7), PS 20 (Figure 3.8), and Pluronic® F68 (Figure 3.9). Hydrophobic interaction is enhanced with increasing temperature, and for each surfactant, accelerated surfactant adsorption in the presence of protein is seen to increase with temperature. Again, this effect is seen to be clearly more pronounced for the polysorbates than for Pluronic® F68. 80 Surface tension (mN/m) 75 70 5 ppm PS 80 5C 5 ppm PS 80 +Lysozyme 5C 5 ppm PS 80 25C 5 ppm PS 80 +Lysozyme 25C 5 ppm PS 80 40C 5 ppm PS 80 +Lysozyme 40C 65 60 55 50 0 50 100 150 200 250 Time (sec) Figure 3.7: Adsorption isotherm (surface tension vs. time) of 5 ppm PS 80 and lysozyme mixture at 5 °C, 25 °C, and 40 °C in 1X PBS. 54 80 Surface tension (mN/m) 75 70 5ppm PS 20 5C 5ppm PS 20 +Lysozyme 5C 5ppm PS 20 25C 5ppm PS 20 +Lysozyme 25C 5ppm PS 20 40C 5ppm PS 20 +Lysozyme 40C 65 60 55 50 0 50 100 150 200 250 Time (sec) Figure 3.8: Adsorption isotherm (surface tension vs. time) of 5 ppm PS 20 and lysozyme mixture at 5 °C, 25 °C, and 40 °C in 1X PBS. 80 Surface tension (mN/m) 75 70 5 ppm F68 5C 5 ppm F68 +Lysozyme 5C 5 ppm F68 25C 5 ppm F68 +Lysozyme 25C 5 ppm F68 40C 5 ppm F68 +Lysozyme 40C 65 60 55 50 0 50 100 150 200 250 Time (sec) Figure 3.9: Adsorption isotherm (surface tension vs. time) of 5 ppm Pluronic ® F68 and lysozyme mixture at 5 °C, 25 °C, and 40 °C in 1X PBS. It is instructive to invoke classical transition-state theory [20] in order to consider the effect of temperature in terms of an equilibrium between the “reactants” in this case (i.e., surfactant aggregates and dissolved protein) and the protein-surfactant activated complex (i.e., the structural intermediate described with reference to Figure 3.1). 55 For this purpose, the initial decrease in surface tension (𝑖𝑛𝑖𝑡𝑖𝑎𝑙 𝑟𝑎𝑡𝑒) at each temperature and surfactant concentration (𝐶) tested can be represented as 𝑖𝑛𝑖𝑡𝑖𝑎𝑙 𝑟𝑎𝑡𝑒 = 𝑘𝑚 ∙ 𝐶 𝑛 , where 𝑘𝑚 is the rate constant characterizing the initial decrease in surface tension and 𝑛 is the order of the reaction. In order to determine the initial rate in each case, surface tension vs. time (𝑡) data were fit to a simple three-parameter exponential decay function 𝑓(𝑡) = 𝑎𝑒 � 𝑏 � 𝑡+𝑐 and initial slopes calculated. The rate constant and reaction order are then easily determined by analysis of plots of 𝑙𝑛 (𝑖𝑛𝑖𝑡𝑖𝑎𝑙 𝑟𝑎𝑡𝑒) vs. 𝑙𝑛 𝐶 for each surfactant at each temperature. The activation energy, 𝐸𝑎 , for adsorption in each case can then be derived from the slope of a plot of ln 𝑘𝑚 vs. 1/𝑇, according to the Arrhenius equation: 𝑙𝑛 𝑘𝑚 = − � 𝐸𝑎 1 ∙ � + 𝑙𝑛 𝐴, 𝑅 𝑇 where 𝑅 is the gas constant, 𝑇 is temperature, and 𝐴 is the Arrhenius pre-exponential factor (Figure 3.10). 56 20 18 ln km F68 F68 + Lysozyme PS 80 PS 80 + Lysozyme PS 20 PS 20 + Lysozyme 10 8 6 4 0.0031 0.0032 0.0033 0.0034 1/T, °C 0.0035 0.0036 0.0037 -1 Figure 3.10: Arrhenius plot of adsorption during initial surface tension depression. The energy of activation, 𝐸𝑎 , was derived from the slope of linear regressions. Considering the effect of temperature in terms of an equilibrium between reactants and an activated complex, the rate constant can be described in thermodynamic terms: 𝑘𝑚 = 𝑘𝑏 𝑇 ℎ ∙𝑒 −∆𝐺� 𝑅𝑇 , where kb and h are Boltzmann’s and Planck’s constants, respectively, and ΔG is the free energy of transition. Thus, since ∆𝐺 = ∆𝐻 − 𝑇 ∙ ∆𝑆, 𝑙𝑛 𝑘𝑚 𝑇 =− ∆𝐻 𝑅 1 ∙ + �𝑙𝑛 𝑇 𝑘𝑏 ℎ + ∆𝑆 𝑅 �. The slope and intercept of plots of 𝑙𝑛 𝑘𝑚 ⁄𝑇 vs. 1⁄𝑇 therefore provide estimates of the enthalpy (ΔH) and entropy (ΔS) of the transition (Figure 3.11). ∆𝐺 can thus be calculated at any desired temperature. Table 3.1 lists the thermodynamic parameters calculated in this fashion for surfactant solutions in the presence and absence of dissolved protein. A reduction in ∆𝐺 was observed in each case upon addition of 57 protein, with the reduction being more pronounced for the polysorbates than for Pluronic® F68. With increases observed in the entropic contribution to free energy (−𝑇 ∙ ∆𝑆) in the presence of protein for each surfactant tested, the entries in Table 3.1 indicate the reduction in ∆𝐺 observed in the presence of protein resulted entirely from a reduction in ΔH. The likely source for this reduction would be an alteration of the van der Waals interactions among the regions on the surfactant molecules mediating their selfassociation. Separation of surfactant monomers from aggregates would disrupt van der Waals interactions and produce an unfavorable enthalpy. Thus based on this analysis we suggest that protein is accelerating the adsorption of the surfactants not by reducing the entropic barrier faced by the surfactant in moving from the aggregate to interface, but by disrupting the surfactant self-association. 16 14 F68 F68 F68++Lysozyme Lysozyme F68 PS PS80 80 PS PS80 80++Lysozyme Lysozyme PS PS20 20 PS PS20 20++Lysozyme Lysozyme ln (km/T) 12 6 4 2 0 0.0031 0.0032 0.0033 0.0034 1/T, °C 0.0035 0.0036 0.0037 -1 Figure 3.11: Determination of thermodynamic components of the activation barrier. The slope and intercept of linear regression provide basis for calculating ∆𝐻 and ∆𝑆, respectively. 58 reaction order Ea(kJ/(mol*K)) delG at 40C (kJ/mole) Enthalpy(H) (kJ/mole) Entropy(S) (J/(K*mole) -TdelS at 40C F68 2.59 17.75 27.97 15.31 -40.47 12.67 F68+Lysozyme 2.65 10.00 26.98 7.56 -62.07 19.43 PS 80 1.64 56.40 54.13 53.96 -0.56 0.18 PS80 + Lysozyme 1.77 34.12 51.50 31.68 -63.35 19.83 PS20 1.75 25.79 52.91 23.35 -94.47 29.57 PS20+Lysozyme 1.81 15.20 51.46 12.75 -123.66 38.71 Table 3.1: Thermodynamic parameters for adsorption of Pluronic® F68, PS 80 and PS 20 in the absence and presence of lysozyme 3.3.4. Effect of protein Lysozyme is structurally stable and highly cationic under the solution conditions used here. Based on the analysis above, we would expect that addition of a more hydrophobic, surface active protein in place of lysozyme would result in greater enhancement of surfactant adsorption than that recorded in the presence of lysozyme. Figures 3.12-3.14 show the initial kinetics of surface tension depression recorded for selected concentrations of PS 80 (Figure 3.12), PS 20 (Figure 3.13), and Pluronic® F68 (Figure 3.14), in the presence and absence of the recombinant protein. At 1 mg/mL, the recombinant protein reduced surface tension to about 54 mN/m after 220 s, while lysozyme had reduced surface tension to only 64 mN/m (at 5 mg/mL) after the same time period. The recombinant protein alone thus displayed more surface tension depression than any of the surfactants at concentrations of 20 ppm and below (Figures 3.2-3.4). Considering only surfactant concentrations where surface tension depression by the surfactant alone is greater than that due to the recombinant protein alone, Figures 3.12 and 3.13 show the adsorption rate was enhanced for each polysorbate by the presence of the recombinant protein at higher concentrations (50 and 100 ppm) than observed for lysozyme. The effect of added protein was less apparent in the case of Pluronic® F68. As described above with reference to lysozyme, these results are consistent with recombinant protein facilitating aggregate disruption, leading either to an increased concentration of 59 surfactant monomers enhancing adsorption (in the case of PS 20 and PS 80), or to the formation of stable, surfactant-protein complexes having little or no effect on surfactant adsorption rate (in the case of Pluronic® F68). Also shown in Figures 3.12-3.14 is the initial kinetics of surface tension depression recorded for each surfactant at 10 ppm, in the presence and absence of the recombinant protein. At 10 ppm, surfactant adsorption is substantially slower than recombinant protein adsorption, but at this surfactant concentration, surfactant-protein mixtures still show greater surface tension depression than either component acting alone (in the case of the polysorbates, Figures 3.12 and 3.13). While we must expect that the surface is dominated by protein in these cases, we hold the enhanced rate recorded is the result of an increased availability of surfactant monomers at the interface owing to protein-mediated disruption of polysorbate aggregates. 75 Surface Tension (mN/m) 70 65 10 ppm PS 80 10 ppm PS 80 + rec.protein 50 ppm PS 80 50 ppm PS 80 + rec.protein 100 ppm PS 80 100 ppm PS 80 + rec.protein rec.protein 60 55 50 45 40 0 50 100 150 200 250 Time (sec) Figure 3.12: Adsorption isotherm (surface tension vs. time) of PS80 and recombinant protein mixture at 25 °C in 10 mM sodium acetate, 5% sorbitol, pH 3.5 60 75 Surface tension (mN/m) 70 65 10 ppm PS 20 10 ppm PS 20 + rec.protein 50 ppm PS 20 50 ppm PS 20 + rec.protein 100 ppm PS 20 100 ppm PS 20 + rec.protein rec.protein 60 55 50 45 40 0 50 100 150 200 250 Time (sec) Figure 3.13: Adsorption isotherm (surface tension vs. time) of PS 20 and recombinant protein mixture at 25 °C in 10 mM sodium acetate, 5% sorbitol, pH 3.5 75 Surface tension (mN/m) 70 65 10 ppm F68 10 ppm F68 + rec.protein 50 ppm F68 50 ppm F68 + rec.protein 100 ppm F68 100 ppm F68 + rec.protein rec.protein 60 55 50 45 40 0 50 100 150 200 250 Time (sec) Figure 3.14. Adsorption isotherm (surface tension vs. time) of Pluronic® F68 and recombinant protein mixture at 25 °C in 10 mM sodium acetate, 5% sorbitol, pH 3.5 3.4 Conclusions We have considered protein effects on surface tension depression by surfactants commonly used in biopharma, in relation to a mechanism for surfactant 61 adsorption governed by the formation of a rate-limiting structural intermediate (i.e., an “activated complex” comprised of surfactant aggregates and protein). A simple thermodynamic analysis indicated the presence of protein caused a reduction in ∆𝑮 for the surfactant adsorption process, with this reduction deriving entirely from a reduction in ∆𝑯. We suggest that protein accelerates the adsorption of these surfactants by disrupting their self associations, releasing surfactant monomers. The increased concentration of surfactant monomers may promote surfactant adsorption, or the formation of stable, surfactant-protein complexes having little or no effect on surfactant adsorption rate. Based on this, we expect that accelerated surfactant adsorption in the presence of protein will occur with surfactants that stabilize protein mainly by their own adsorption at interfaces, and we expect the absence of accelerated surfactant adsorption will occur with surfactants that form stable surfactant-protein associations. This expectation is currently being tested with the proteins and surfactants used here, by revealing the dominant mode of surfactant-mediated stabilization of protein at solid surfaces, and will constitute the subject of a future report. 62 3.5 References [1] J. Carpenter, B. Cherney, A. Lubinecki, S. Ma, E. Marszal, A. Mire-Sluis, T. Nikolai, J. Novak, J. Ragheb, and J. Simak, Meeting report on protein particles and immunogenicity of therapeutic proteins: filling in the gaps in risk evaluation and mitigation. Biologicals 38 (2010) 602-11. [2] W. Wang, S. Nema, and D. Teagarden, Protein aggregation--pathways and influencing factors. Int J Pharm 390 (2010) 89-99. [3] J.S. Philo, and T. Arakawa, Mechanisms of protein aggregation. Curr Pharm Biotechnol 10 (2009) 348-51. [4] E.Y. Chi, S. Krishnan, T.W. Randolph, and J.F. Carpenter, Physical stability of proteins in aqueous solution: mechanism and driving forces in nonnative protein aggregation. Pharm Res 20 (2003) 1325-36. [5] J.S. Bee, T.W. Randolph, J.F. Carpenter, S.M. Bishop, and M.N. Dimitrova, Effects of surfaces and leachables on the stability of biopharmaceuticals. J Pharm Sci (2011). [6] O. Joshi, J. McGuire, and D.Q. Wang, Adsorption and function of recombinant factor VIII at solid-water interfaces in the presence of Tween-80. J Pharm Sci 97 (2008) 4741-55. [7] D.K. Chou, R. Krishnamurthy, T.W. Randolph, J.F. Carpenter, and M.C. Manning, Effects of Tween 20 and Tween 80 on the stability of Albutropin during agitation. J Pharm Sci 94 (2005) 1368-81. [8] O. Joshi, L. Chu, J. McGuire, and D.Q. Wang, Adsorption and function of recombinant Factor VIII at the air-water interface in the presence of Tween 80. J Pharm Sci 98 (2009) 3099-107. [9] H.J. Lee, A. McAuley, K.F. Schilke, and J. McGuire, Molecular origins of surfactant-mediated stabilization of protein drugs. Adv Drug Deliv Rev 63 (2011) 116071. [10] V. Schram, and S.B. Hall, Thermodynamic effects of the hydrophobic surfactant proteins on the early adsorption of pulmonary surfactant. Biophys J 81 (2001) 1536-46. [11] E.D. Goddard, N.J. Turro, P.L. Kuo, and K.P. Ananthapadmanabhan, Fluorescence probes for critical micelle concentration determination. Langmuir 1 (1985) 352-5. [12] S.R. Croy, and G.S. Kwon, Polysorbate 80 and Cremophor EL micelles deaggregate and solubilize nystatin at the core-corona interface. J Pharm Sci 94 (2005) 2345-54. [13] B.A. Kerwin, Polysorbates 20 and 80 used in the formulation of protein biotherapeutics: structure and degradation pathways. J Pharm Sci 97 (2008) 2924-35. [14] P. Bahadur, P. Li, M. Almgren, and W. Brown, Effect of potassium fluoride on the micellar behavior of Pluronic F-68 in aqueous solution. Langmuir 8 (1992) 19031907. [15] H.W. Tsui, Y.H. Hsu, J.H. Wang, and L.J. Chen, Novel behavior of heat of micellization of pluronics PLURONIC ® F68 and F88 in aqueous solutions. Langmuir 24 (2008) 13858-62. [16] Y. Li, M. Bao, Z. Wang, H. Zhang, and G. Xu, Aggregation behavior and complex structure between triblock copolymer and anionic surfactants. Journal of Molecular Structure 985 (2011) 391-396. 63 [17] S. Borbély, and J.S. Pedersen, Temperature-induced aggregation in aqueous solutions of pluronic PLURONIC ® F68 triblock copolymer containing small amount of o-xylene. Physica B: Condensed Matter 276–278 (2000) 363-364. [18] Y. Levin, A.P. dos Santos, and A. Diehl, Ions at the air-water interface: an end to a hundred-year-old mystery? Phys Rev Lett 103 (2009) 257802. [19] K.R. Acharya, S.C. Bhattacharyya, and S.P. Moulik, Effects of carbohydrates on the solution properties of surfactants and dye–micelle complexation. Journal of Photochemistry and Photobiology A: Chemistry 122 (1999) 47-52. [20] O. Levenspiel, Chemical reaction engineering, Wiley, New York, 1999. 64 DOMINANT MODE OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN AT SOLID SURFACES Hyo Jin Lee,a,b Arnold McAuley,b and Joseph McGuirea,* a School of Chemical, Biological and Environmental Engineering, Oregon State University, Corvallis, OR 97331 b Proces and Product Development Department Amgen Inc., Thousand Oaks, CA 91320 * Corresponding Author: Joseph McGuire School of Chemical, Biological and Environmental Engineering Oregon State University 103 Gleeson Hall Corvallis, OR 97331-2702 Tel: 541-737-6306 Fax: 541-737-4600 Email: mcguirej@engr.orst.edu 65 CHAPTER 4 DOMINANT MODE OF SURFACTANT-MEDIATED STABILIZATION OF PROTEIN AT SOLID SURFACES Abstract In earlier work we considered protein effects on surface tension depression by the surfactants polysorbate 20, polysorbate 80, and Pluronic® F68, in relation to a mechanism for surfactant adsorption governed by the formation of a ratelimiting structural intermediate Based on results of that work, we proposed that accelerated surfactant adsorption in the presence of protein (observed with PS 20 and PS 80) will occur with surfactants that stabilize protein mainly by their own adsorption at interfaces, and that the absence of accelerated surfactant adsorption (observed with F68) will occur with surfactants that form stable surfactant-protein associations. Optical waveguide lightmode spectroscopy was used to test this expectation. Adsorption kinetics were recorded for surfactants (PS 20, PS 80, or F68) and protein (lysozyme or Amgen recombinant protein) at a hydrophilic solid (SiO2-TiO2) surface. Experiments were performed in sequential and competitive adsorption modes, enabling the adsorption kinetic patterns to be interpreted in a fashion revealing the dominant mode of surfactantmediated stabilization of protein in each case. Kinetic results confirmed predictions based on our earlier quantitative analysis of protein effects on surface tension depression by surfactants. In particular, PS 20 and PS 80 are able to inhibit protein adsorption only by their preferential location at the interface, and not by formation of less surface active, protein-surfactant complexes. On the other hand, F68 clearly is able to inhibit protein adsorption by formation of protein-surfactant complexes, and not by its preferential location at the interface. 66 4.1 Introduction In the previous chapter we considered protein effects on surface tension depression by the surfactants polysorbate 20, polysorbate 80, and Pluronic® F68, in relation to a mechanism for surfactant adsorption governed by the formation of a ratelimiting structural intermediate (i.e., an “activated complex” comprised of surfactant aggregates and protein). A simple thermodynamic analysis indicated the presence of protein caused a reduction in ∆𝑮 for the surfactant adsorption process (and consequent acceleration of surface tension depression), with this reduction deriving entirely from a reduction in ∆𝑯. We suggested that in cases where protein accelerated the adsorption of the surfactants (observed with PS 20 and PS 80); it did so by disrupting their self associations, releasing surfactant monomers. The increased concentration of surfactant monomers then promoted surfactant adsorption. The absence of accelerated surface tension depression (observed with F68) was attributed to the newly released monomers participating in the formation of stable, surfactant-protein complexes thus having little or no effect on surfactant adsorption rate. Based on that work, we would expect that accelerated surfactant adsorption in the presence of protein will occur with surfactants that stabilize protein mainly by their own adsorption at interfaces, and would expect the absence of accelerated surfactant adsorption will occur with surfactants that form stable surfactant-protein associations. In this paper we use optical waveguide lightmode spectroscopy to test this expectation. Adsorption kinetics were recorded for surfactants (PS 20, PS 80, or F68) and protein (lysozyme or Amgen recombinant protein) at a hydrophilic solid (SiO2-TiO2) surface. The hydrophilic surface was chosen to enable study of adsorption kinetics in cases for which protein-surface affinity is high in relation to (nonionic) surfactant-surface 67 affinity. Experiments were performed in sequential and competitive adsorption modes, enabling the adsorption kinetic patterns to be interpreted in a fashion revealing the dominant mode of surfactant-mediated stabilization of protein in each case. 4.2 Material and Methods 4.2.1. Materials and sample preparation The standard sensor chips used for monitoring adsorption kinetics are obtained from MicroVacuum Inc (Budapest, Hungary). It is made by depositing a thin layer of Si0.25Ti0.75O2 by a sol-gel process to a glass chip. An optical waveguide grating, which is approximately 20 nm deep, sinusoidal, and has 2400 lines/mm, is embossed to the surface of the chip [4]. The sensor chips were used without further modification and were soaked in the desired buffer overnight to enable equilibration of the waveguide film with the cover medium. The proteins used were lyophilized chicken egg white lysozyme (Sigma Aldrich, Saint Louis, MO) and a recombinant protein provided by Amgen. Lysozyme was dissolved in Dulbecco’s phosphate buffered saline (1X PBS, Invitrogen, Carlsbad, CA) at 0.5 mg/mL , which equals to 0.073mM and was filtered using a 0.2 um PVDF filter (Millipore, Billerica, MA). The recombinant protein is approximately 18.8 kDa that consists of 174 amino acids with 2 disulfide bonds. It was provided in 10 mM sodium acetate, 5% sorbitol at pH 4.0 at 3 mg/mL and was diluted to 0.4 mg/mL, which equals to 0.073mM and filtered using a 0.2um PVDF filter before use. Both protein concentrations were measured with a UV spectrophotometer (Cary 50, Varian, Inc.) using 0.66 mg/ mL cm as extinction coefficient for lysozyme and 0.78 for the recombinant protein. 68 The surfactants studied were Poloxamer 188 (Pluronic ® F68, BASF, obtained from Sigma Aldrich, Saint Louis, MO) polysorbate 80 (PS 80, Croda, East Yorkshire, England) and polysorbate 20 (PS 20, Croda, East Yorkshire, England). Stock solutions at high concentrations of the surfactants were made in 1X PBS and in 10 mM sodium acetate, 5% sorbitol at pH 4.0 buffers and diluted to the respective buffers to have a molar ratio of 1:10 and 1:100 with the proteins evaluated. Buffer salts and sorbitol were from obtained from JT Baker (Philipsburg, NJ). To study the effect of salt, the recombinant protein was dialyzed in 10mM sodium acetate, 250mM sodium chloride, 5% sorbitol, at pH 4.0 using a Slide-Alyzer (10KDa MW cutoff, Thermo Fisher Scientific, Rockford, IL). Concentration was measured with a UV spectrophotometer (Cary 50, Varian, Inc.) and diluted to 0.4mg/mL. 4.2.2. Evaluation of adsorption kinetics The adsorption kinetics was recorded in-situ by Optical Waveguide Lightmode Spectroscopy (OWLS, MicroVacuum Inc. Hungary). The basic principle of operation is when linearly polarized HeNe (632.8 nm) laser is coupled by a diffraction grating into a waveguide layer. At two distinct angles, light is propagated by total internal reflection along the waveguide which is defined as incoupling. The properties of the covering media determine the specific incident angle necessary for the light to propagate, which is defined as the incoupling angle. From the angle and mathematical optics models, the refractive index (𝑑𝐴 ) and thickness (𝑛𝐴 ) of the overlayer can be calculated. These can then be used to estimate the immobilized mass (Γ) of the adsorbed overlayer [5]: 𝑑𝑛 −1 𝛤 = 𝑑𝐴 (𝑛𝐴 − 𝑛𝐶 ) � � 𝑑𝑐 69 where 𝐶 is the solution concentration of the protein, 𝑛𝐶 is the bulk solution refractive 𝑑 index, and 𝑑𝑛 is the refractive index increment and is usually 0.182 cm3/g for proteins in 𝑐 solution which was used consistently throughout all measurements. The sensor chips that were soaked in the desired buffer overnight are placed in a flow cell. After aligning the sensor chip properly and securing it tightly to prevent any buffer leaks, the syringe pump is turned on to flow the desired buffer of interest through the system at 100 µL/min. After establishing a stable baseline, 100 µL of protein alone, surfactant alone, or protein+ surfactant mixture, which will be described in detail in the following subsections, was injected using a Hamilton syringe via valve diverter while the syringe pump was turned off. Adsorption was monitored for 30 minutes. The sample was then rinsed with the same buffer at 100 µL /min for 15 minutes. 4.2.2.1. Surfactant+protein co-adsorption Protein concentration was kept constant and the amount of surfactant was varied to obtain a protein to surfactant molar ratio of 1:10 and 1:100. Final concentration of poloxamer (Pluronic® F68) was at 600ppm and 6000ppm and polysorbate 80 (PS 80) and 20 (PS20) were studied at 90ppm and 900ppm. After stable baselines were achieved, the protein and surfactant mixtures, which were incubated for at least 1 hour, were introduced and monitored for 30 minutes followed by a rinsing step just by buffer for 15 minutes. 4.2.2.2. Surfactant+protein sequential adsorption After running buffer achieved stable baseline signals, first Pluronic® F68 at 600 ppm was introduced and monitored for 30 minutes followed by a buffer rinsing step for 15 minutes. A rinsing step was added to remove any surfactant in the bulk or 70 reversibly bound surfactant to the sensor. Then, the protein of interest was added and monitored for 30 minutes followed by a buffer rinsing step for another 15 minutes. 4.2.2.3. Surfactant+protein co-adsorption pre-coated with surfactant After buffer stabilization was achieved, 90ppm PS 80 or PS 20 was introduced and monitored for 30 minutes. To maintain same concentration of surfactant throughout the experiment, no rinse step was involved. After 30 minutes, protein mixtures with 90ppm PS 80 or PS 20 was introduced and monitored for another 30 minutes followed by a buffer rinse for 15 minutes. 4.3 Results and Discussion 4.3.1. Surfactant+lysozyme co-adsorption The adsorption kinetics recorded for surfactant solutions in the absence of protein are presented in Figure 4.1. Both PS 80 and PS 20 displayed higher affinity for the surface than Pluronic® F68 at each concentration tested, with each surfactant adsorbing in a fashion not entirely elutable by surfactant-free buffer. The adsorption kinetics of lysozyme in PBS in the absence and presence of PS 80 and PS 20 are shown in Figure 4.2. While the presence of surfactant altered the adsorption kinetics to some extent, upon elution the kinetic patterns were highly similar to that recorded for lysozyme in the absence of surfactant, in each case. Joshi et al. found polysorbate 80 to have no measurable effect on the adsorption of a human recombinant Factor VIII at a hydrophilic silica surface [6]. While each polysorbate showed some affinity for the surface used in the present experiments, it is possible that these adsorbed layers are dominated by lysozyme after elution. 71 Adsorbed amount (ng/cm2) 200 90 ppm PS 80 900 ppm PS 80 90 ppm PS 20 900 ppm PS 20 600 ppm F68 6000 ppm F68 150 100 50 0 0 10 20 30 40 50 Time (min) Figure 4.1: Adsorption kinetics of 90 and 900 ppm PS 80, 90 and 900 ppm PS 20, and 600 and 6000 ppm Pluronic® F68 in PBS 72 200 Adsorbed amount(ng/cm2) 0 ppm 900 ppm PS 80 900 ppm PS 20 150 100 50 0 0 10 20 30 40 50 Time (min) Figure 4.2: Adsorption kinetics of lysozyme in PBS in the presence of 0 ppm, 900 ppm PS 80 and 900 ppm PS 20 The adsorption kinetics of lysozyme in PBS in the presence of 0 ppm, 600ppm and 6000ppm Pluronic® F68 are shown in Figure 4.3. In contrast to the patterns observed with the polysorbates, the amount of adsorption after the elution step is reduced appreciably in the presence of F68, to a level near that expected for F68 in the absence of protein. This could be due to F68 locating at the interface more rapidly than lysozyme, thus inhibiting lysozyme adsorption. On the other hand, the F68 triblocks may have associated with lysozyme in solution to form complexes of low adsorption affinity. As shown in Figure 4.1 however, F68 itself shows very low affinity for the surface, and these results likely suggest that the reduction in protein adsorption is owing to complex formation and not to F68 location at the surface. This can be tested by the sequential 73 introduction of F68 and lysozyme to the surface. 200 Adsorbed amount (ng/cm2) 0 ppm 600 ppm F68 6000 ppm F68 150 100 50 0 0 10 20 30 40 50 Time (min) Figure 4.3: Adsorption kinetics of lysozyme in PBS in the presence of 0 ppm, 600 ppm, and 6000 ppm Pluronic® F68 4.3.2. Pluronic® F68 + lysozyme sequential adsorption Figure 4.4 shows results of a sequential adsorption experiment in which introduction of F68 to the surface is followed by introduction of lysozyme. This intermediate rinse step at 30 min removes any loosely bound F68 at the interface as well as any F68 present in solution. (Data recorded for lysozyme with no F68, from Figure 4.2, was superimposed on Figure 4.4 beginning at 45 min for comparison.) As shown clearly in Figure 4.4, the presence of surface-bound F68 did not prevent lysozyme adsorption. Upon addition of lysozyme at 45 min, the adsorption kinetics were somewhat slower than that recorded without the F68 “pre-coated” on the surface, possibly owing to 74 displacement of adsorbed F68 by lysozyme. The elution patterns are highly similar in each case. These observations strongly support the thought that F68 does not inhibit lysozyme adsorption by its preferential location to the interface. Rather, F68 inhibits lysozyme adsorption by formation of less surface active, protein-surfactant complexes. 200 0 ppm 600 ppm F68 Adsorbed amount (ng/cm2) Lysozyme 150 100 Rinse Rinse 50 0 0 20 40 60 80 100 Time (min) Figure 4.4: Adsorption kinetics of 600 ppm Pluronic® F68 followed by buffer elution and the introduction of lysozyme in PBS. 4.3.3. Polysorbates +lysozyme co-adsorption pre-coated with polysorbates Figure 4.5 shows polysorbate adsorption followed by the addition of lysozyme + polysorbate, followed by elution with PBS buffer. (Data recorded for lysozyme with no polysorbate, from Figure 4.2, was superimposed on Figure 4.5 beginning at 30 min for comparison.) PS 80 and PS 20 displayed appreciable adsorption upon introduction, and were not eluted. Upon introduction of protein in the presence of 75 the PS 80, the adsorption kinetics were somewhat slower than that recorded without PS 80 “pre-coated” on the surface, likely owing to displacement of adsorbed PS 80 by lysozyme. No appreciable change in kinetics was observed in the case of PS 20 + lysozyme in relation to lysozyme alone, consistent with PS 80 showing a higher surface affinity than PS 20. Finally, as in Figure 4.2, upon elution the kinetic patterns were highly similar to that recorded for lysozyme in the absence of surfactant, in each case. 200 0 ppm 90 ppm PS80 90 ppm PS 20 Adsorbed amount (ng/cm2) Lysozyme + polysorbates 150 100 Rinse 50 0 0 20 40 60 80 Time(min) Figure 4.5: 90 ppm polysorbates adsorption followed by 90 ppm polysorbates + lysozyme co-adsorption in PBS 4.3.4. Surfactants+ recombinant protein co-adsorption The adsorption kinetics recorded for only surfactant solutions in 10 mM sodium acetate, 5% sorbitol, pH 4.0 are presented in Figure 4.6. Both PS 80 and PS 20 displayed higher affinity for the surface than Pluronic® F68 at each concentration tested, 76 with each surfactant adsorbing in a fashion not entirely elutable by surfactant-free buffer. Also, all of the surfactants displayed higher affinity in 10 mM sodium acetate, 5% sorbitol, pH 4.0 compared to when in PBS (Figure 4.1). The adsorption kinetics of recombinant protein in 10 mM sodium acetate, 5% sorbitol, pH 4.0 in the absence and presence of PS 80, PS 20 and F68 are shown in Figure 4.7. Only the highest surfactant concentrations are shown since the lower surfactant concentrations displayed similar adsorption and elution behavior. All of the surfactants are seen to have a slight effect in reducing the amount of protein on the surface upon rinsing. However, differences in the elution patterns are observed when comparing lysozyme to recombinant protein in the presence of surfactants. In the presence of F68, the amount of adsorption of the recombinant protein after the elution step did not decrease significantly as observed with lysozyme. However, in the presence of polysorbates, more effect was seen in the elution step for the recombinant protein than for lysozyme which had almost none. In a previous study, recombinant protein is shown to display more surface activity than lysozyme at the air-water interface. Consequently, more adsorption and less rinsing of the recombinant protein is seen when compared to lysozyme at the same surface. The effect of surfactants on the adsorption kinetics could be due to the recombinant protein being more surface active than lysozyme. Also, the surfactants, especially the polysorbates showed higher affinity in 10 mM sodium acetate, 5% sorbitol, at pH 4.0 than in PBS. This could be another factor for differences shown for the recombinant protein. 77 250 90 ppm PS 80 900 ppm PS 80 90 ppm PS 20 900 ppm PS 20 600 ppm F68 6000 ppm F68 Adsorbed amount (ng/cm2) 200 150 100 50 0 0 10 20 30 40 50 Time (min) Figure 4.6: Adsorption kinetics of 90 and 900 ppm PS 80, 90 and 900 ppm PS 20, and 600 and 6000ppm Pluronic® F68 in 10 mM sodium acetate, 5 % sorbitol, at pH 4.0 78 250 Adsorbed amount (ng/cm2) 200 150 100 0 ppm 6000 ppm F68 900 ppm PS 80 900 ppm PS 20 50 0 0 10 20 30 40 50 Time (min) Figure 4.7: Adsorption kinetics of recombinant protein in 10 mM sodium acetate, 5% sorbitol, pH 4.0 in in the presence of 0 ppm, 900 ppm PS 80, 900 ppm PS 20, and 6000ppm Pluronic® F68 4.3.5. Polysorbates+ recombinant protein co-adsorption pre-coated with polysorbates Figure 4.8 shows polysorbate adsorption followed by the addition of recombinant protein + polysorbate, followed by elution with 10 mM sodium acetate, 5% sorbitol, pH 4.0 buffer. (Data recorded for recombinant with no polysorbate, from Figure 4.7, was superimposed on Figure 4.8 beginning at 30 min for comparison.) PS 80 and PS 20 displayed appreciable adsorption upon introduction, and were not eluted. Upon introduction of protein in the presence of the PS 80, the adsorption kinetics were somewhat slower than that recorded without PS 80 “pre-coated” on the surface, likely owing to displacement of adsorbed PS 80 by recombinant protein. No appreciable change in kinetics was observed in the case of PS 20 + recombinant protein in relation to 79 recombinant protein alone, consistent with PS 80 showing a higher surface affinity than PS 20. Due to the higher affinity of the polysorbates at the surface, the amount of adsorption after the elution step is reduced in the presence of polysorbates, more significant for PS 80 than for PS 20. 250 Recombinant protein +Polysorbates Adsorbed amount (ng/cm2) 200 150 Rinse 100 50 0 ppm 90 ppm PS 80 90 ppm PS 20 0 0 20 40 60 Time (min) Figure 4.8: 90 ppm polysorbates adsorption followed by 90 ppm polysorbates + recombinant protein co-adsorption in 10 mM sodium acetate, 5 % sorbitol, pH 4.0 4.3.6. Effect of salt on Pluronic® F68 +recombinant protein co-adsorption and sequential adsorption Our previous paper has shown that a mechanism for surfactant adsorption may be governed by the formation of a rate-limiting structural intermediate (i.e., an “activated complex”) between surfactant aggregates and protein [3]. While Pluronic® F68 is not expected to form micelles in water except at very high 80 80 concentrations[7-10]; it is known to exhibit greater aggregation and micellization in salt solutions[11]. Since Pluronic® F68 was not as effective in reducing the recombinant protein adsorption compared to lysozyme; addition of 250 mM in 10 mM sodium acetate, 5 % sorbitol, at pH 4.0 was studied to verify the role of surfactant aggregates. As shown in Figure 4.9, the amount of recombinant protein adsorbed reduced when the ionic strength was increased. The presence of salt at higher concentrations effectively shields the charges on the surface and on the protein which is well documented. The adsorption kinetics of recombinant protein in 10 mM sodium acetate, 5% sorbitol, 250 mM sodium chloride, at pH 4.0 in the presence of 0 ppm, 600ppm and 6000ppm Pluronic® F68 are shown in Figure 4.10. Reduction of recombinant protein adsorption with Pluronic® F68 in the elution step is more significant in the presence of salt than without salt as shown in Figure 4.7. The addition of 250 mM sodium chloride may have induced Pluronic® F68 aggregation which associated with recombinant protein more, thus forming lower surface affinity complexes. As a result, less adsorption is observed for recombinant protein in the presence of Pluronic® F68 in 250mM sodium chloride. Figure 4.11 shows results of a sequential adsorption experiment in which introduction of F68 to the surface is followed by introduction of recombinant protein. This intermediate rinse step at 30 min removes any loosely bound F68 at the interface as well as any F68 present in solution. (Data recorded for recombinant protein with no F68, from Figure 4.10, was superimposed on Figure 4.11 beginning at 45 min for comparison.) As shown clearly in Figure 4.11, the presence of surface-bound F68 did not prevent recombinant protein adsorption. Upon addition of recombinant protein at 45 min, the adsorption kinetics were similar with that recorded without the F68 “pre-coated” on the surface. The elution patterns are highly similar in each case. These observations strongly 81 support the thought that F68 does not inhibit recombinant protein adsorption by its preferential location to the interface. Rather, in the presence of salt, F68 inhibits recombinant protein by formation of less surface active, protein-surfactant complexes. 250 Adsorbed amount (ng/cm2) 200 150 100 50 0 mM NaCl 250 mM NaCl 0 0 10 20 30 40 50 Time (min) Figure 4.9: Effect of salt concentration on the adsorption of recombinant protein in 10 mM sodium acetate, 5 % sorbitol, at pH 4.0 82 250 0 ppm + 250 mM NaCl 600 ppm F68 + 250 mM NaCl 6000 ppm F68 + 250 mM NaCl Adsorbed amount (ng/cm2) 200 150 100 50 0 0 10 20 30 40 50 Time (min) Figure 4.10: Adsorption kinetics of recombinant protein in 10 mM sodium acetate, 5 % sorbitol, 250 mM sodium chloride, at pH 4.0 in the presence of 0 ppm, 600ppm, and 6000ppm Pluronic® F68 83 250 Rinse Recombinant protein Adsorbed amount (ng/cm2) 200 150 100 Rinse 50 0 ppm + 250 mM NaCl 600 ppm F68 + 250 mM NaCl 0 0 20 40 60 80 100 Time (min) Figure 4.11: Adsorption kinetics of 600ppm Pluronic® F68 followed by buffer elution and the introduction of recombinant protein in 10 mM sodium acetate, 5 % sorbitol, 250 mM sodium chloride, at pH 4.0. 4.4 Conclusions To determine the dominant mode of surfactant-mediated stabilization of protein, lysozyme and a more surface active recombinant protein was studied in the presence of Pluronic® F68, PS 80 and PS 20 at solid surfaces. Pluronic® F68 was effective in reducing protein adsorption especially in the presence of salt by forming lower surface affinity complexes. PS 80 and PS 20 were not effective in preventing lysozyme adsorption and only slight reductions were observed for the recombinant protein adsorption. Even though polysorbates were observed to adsorb at the surface, protein was able to replace all or most of the adsorbed surfactant owing to the weak polysorbate-surface interaction. 84 4.5 References [1] J. Carpenter, B. Cherney, A. Lubinecki, S. Ma, E. Marszal, A. Mire-Sluis, T. Nikolai, J. Novak, J. Ragheb, and J. Simak, Meeting report on protein particles and immunogenicity of therapeutic proteins: filling in the gaps in risk evaluation and mitigation. Biologicals 38 (2010) 602-11. [2] J.S. Bee, T.W. Randolph, J.F. Carpenter, S.M. Bishop, and M.N. Dimitrova, Effects of surfaces and leachables on the stability of biopharmaceuticals. J Pharm Sci (2011). [3] H.J. Lee, A. McAuley, K.F. Schilke, and J. McGuire, Molecular origins of surfactant-mediated stabilization of protein drugs. Adv Drug Deliv Rev 63 (2011) 116071. [4] A. Szekacs, N. Adanyi, I. Szekacs, K. Majer-Baranyi, and I. Szendro, Optical waveguide light-mode spectroscopy immunosensors for environmental monitoring. Appl Opt 48 (2009) B151-8. [5] J.A. De Feijter, J. Benjamins, and F.A. Veer, Ellipsometry as a tool to study the adsorption behavior of synthetic and biopolymers at the air–water interface. Biopolymers 17 (1978) 1759-1772. [6] O. Joshi, J. McGuire, and D.Q. Wang, Adsorption and function of recombinant factor VIII at solid–water interfaces in the presence of Tween-80. J Pharm Sci 97 (2008) 4741-4755. [7] P. Bahadur, P. Li, M. Almgren, and W. Brown, Effect of potassium fluoride on the micellar behavior of Pluronic F-68 in aqueous solution. Langmuir 8 (1992) 19031907. [8] H.-W. Tsui, Y.-H. Hsu, J.-H. Wang, and L.-J. Chen, Novel Behavior of Heat of Micellization of Pluronics F68 and F88 in Aqueous Solutions. Langmuir 24 (2008) 13858-13862. [9] Y. Li, M. Bao, Z. Wang, H. Zhang, and G. Xu, Aggregation behavior and complex structure between triblock copolymer and anionic surfactants. Journal of Molecular Structure 985 (2011) 391-396. [10] S. Borbely, and J.S. Pedersen, Temperature-induced aggregation in aqueous solutions of pluronic F68 triblock copolymer containing small amount of o-xylene. Physica B: Condensed Matter 276-278 (2000) 363-364. [11] K. Patel, B. Bharatiya, Y. Kadam, and P. Bahadur, Micellization and Clouding Behavior of EO–PO Block Copolymer in Aqueous Salt Solutions. Journal of Surfactants and Detergents 13 89-95. 85 CHAPTER 5 GENERAL CONCLUSION A simple thermodynamic analysis indicated the presence of protein caused a reduction in ∆𝑮 for the surfactant adsorption process, with this reduction deriving entirely from a reduction in ∆𝑯. We suggest that protein accelerates the adsorption of these surfactants by disrupting their self associations, releasing surfactant monomers. The increased concentration of surfactant monomers may promote surfactant adsorption, or the formation of stable, surfactant-protein complexes having little or no effect on surfactant adsorption rate. Based on this, we expect that accelerated surfactant adsorption in the presence of protein will occur with surfactants that stabilize protein mainly by their own adsorption at interfaces, and we expect the absence of accelerated surfactant adsorption will occur with surfactants that form stable surfactant-protein associations. Pluronic® F68 was shown to be effective in reducing protein adsorption especially in the presence of salt by forming lower surface affinity complexes. PS 80 and PS 20 were not effective in preventing lysozyme adsorption and only slight reductions were observed for the recombinant protein adsorption. Even though polysorbates were observed to adsorb at the surface, the protein was able to replace all or most of the adsorbed surfactant owing to the weak polysorbate-surface interaction. In summary PS 20 and PS 80 are able to inhibit protein adsorption only by their preferential location at the interface, and not by formation of less surface active, protein-surfactant complexes. On the other hand, F68 clearly is able to inhibit protein adsorption by formation of proteinsurfactant complexes, and not by its preferential location at the interface. 86 Chapters 2-4 present how surfactants interact with interfaces and proteins (particularly the preferential location of a surfactant at an interface, or its association with protein in solution), and can provide guidance in selecting surfactants and excipients to reduce protein losses in a given application. 87 BIBLIOGRAPHY A. Gigout, M.D. Buschmann, M. Jolicoeur, The fate of Pluronic® F-68 in chondrocytes and CHO cells, Biotechnol. 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