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	Preface	V
1	Efficient and Reliable Production of Pharmaceuticals in Alfalfa MARC-ANDRE D’AOUST, PATRICE LEROUGE, URSULA BUSSE, PIERRE BILODEAU, SONIA TRÉPANIER, VÉRONIQUE GOMORD, LOIC FAYE and LOUIS-PHILIPPE VÉZINA	1
1.1	Introduction	1
1.2	Alfalfa-specific Expression Cassettes	2
1.3	Alfalfa Transformation Methods	3
1.4	Characteristics of Alfalfa-derived Pharmaceuticals	6
1.5	Industrial Production of Recombinant Proteins in Alfalfa	9
1.5.1	Ramping Up Alfalfa Biomass	9
1.5.2	Alfalfa Harvest, and Recovery of Recombinant Molecules	10
1.6	Conclusions	11
	References	11
2	Foreign Protein Expression Using Plant Cell Suspension and Hairy Root Cultures FIONA S. SHADWICK and PAULINE M. DORAN	13
2.1	Foreign Protein Production Systems	13
2.2	Production of Foreign Proteins Using Plant Tissue Culture	14
2.2.1	Suspended Cell Cultures	15
2.2.2	Hairy Root Cultures	20
2.2.3	Shooty Teratoma Cultures	20
2.2.4	Scale-up Considerations for Different Forms of Plant Tissue Culture	21
2.3	Strategies for Improving Foreign Protein Accumulation and Product Recovery in Plant Tissue Culture	22
2.3.1	Expression Systems	22
2.3.1.1	Modifications to Existing Expression Constructs	22
2.3.1.2	Transient Expression Using Viral Vectors	23
2.3.2	Secretion of Foreign Proteins	25
2.3.3	Foreign Protein Stability	26
2.3.3.1	Stability Inside the Cells	26
2.3.3.2	Stability Outside the Cells	26
2.3.3.3	Medium Additives	28
2.3.3.4	Medium Properties	31
2.3.4	Bioprocess Developments	33
2.3.4.1	Product Recovery from the Medium	33
2.3.4.2	Oxygen Transfer and Dissolved Oxygen Concentration	33
2.4	Conclusions	34
	References	34
3	Novel Sprouting Technology for Recombinant Protein Production KIMMO KOIVU	37
3.1	Introduction	37
3.2	Biology of Sprouting	38
3.2.1	Structure and Content of Dicotyledonous and Monocotyledonous Seeds	38
3.2.2	Germination	39
3.2.3	The Sprout	40
3.2.4	Rubisco Synthesis	40
3.2.5	Rubisco Promoters	41
3.2.6	Inhibition of Endogenous Gene Expression	42
3.3	Expression Cassette Design	43
3.4	Sprouting Equipment	44
3.5	Sprouting Conditions	45
3.5.1	Sterilization	46
3.5.2	Sprouting Time and Temperature	46
3.5.3	Light	47
3.5.4	Inhibition of Endogenous Gene Expression	47
3.5.5	Growth Regulators	49
3.5.6	Nitrogen Fertilizer	49
3.5.7	Seed Production	49
3.6	Yield Estimates and Benefits of Sprouting Technology in Protein Production	50
3.6.1	Yield Estimates	50
3.6.2	Quality and Environmental Aspects	52
	References	53
4	Monocot Expression Systems for Molecular Farming PAUL CHRISTOU, EVA STOGER and RICHARD M. TWYMAN	55
4.1	Introduction	55
4.2	Cereal Production Crops	56
4.3	Technical Aspects of Molecular Farming in Cereals	57
4.3.1	Cereal Transformation	57
4.3.2	Expression Construct Design	59
4.3.3	Production Considerations for Cereals	61
4.4	Examples of Recombinant Proteins Produced in Cereals	61
4.4.1	ProdiGene and Maize	62
4.4.2	Recombinant Proteins Expressed in Rice	63
4.4.3	Recombinant Proteins Produced in Wheat	64
4.4.4	Recombinant Proteins Produced in Barley	64
4.5	Conclusions	64
	References	65
5	The Field Evaluation of Transgenic Crops Engineered to Produce Recombinant Proteins JIM BRANDLE	69
5.1	Introduction	69
5.2	Regulation of Field-testing	69
5.3	Design of Field Trials	73
5.4	Results of Field Trials	74
	References	75
6	Plant Viral Expression Vectors: History and New Developments VIDADI YUSIBOV and SHAILAJA RABINDRAN	77
6.1	Introduction	77
6.2	Plant RNA Viruses as Expression Vectors	78
6.2.1	Tobacco mosaic virus (TMV)	80
6.2.2	Potato virus X (PVX)	80
6.2.3	Cowpea mosaic virus (CPMV)	81
6.2.4	Alfalfa mosaic virus (AlMV)	81
6.3	Biological Activity of Target Molecules	81
6.4	Efficacy of Plant Virus-produced Antigens	83
6.4.1	Vaccine Antigens	83
6.4.2	Particle-based Vaccine Antigen Delivery	84
6.4.3	Other Uses of Plant Virus Particles	86
6.5	Plant Viruses as Gene Function Discovery Tools	87
6.6	New Approaches to the Development of Viral Vectors	87
6.7	Conclusion	88
	References	89
7	Production of Pharmaceutical Proteins in Plants and Plant Cell Suspension Cultures ANDREAS SCHIERMEYER, SIMONE DORFMÜLLER and HELGA SCHINKEL	91
7.1	Introduction	91
7.2	Plant Species Used for Molecular Farming	92
7.3	Cell Culture as an Alternative Expression System to Whole Plants	99
7.4	From Gene to Functional Protein: Processing Steps in Plants	102
7.5	Case Studies of Improved Protein Yields	104
7.6	Downstream Processing	105
7.7	Market Potential of Plant-derived Pharmaceuticals	106
7.8	Containment Strategies for Molecular Farming	107
7.9	Concluding Remarks	108
	References	109
8	Chloroplast Derived Antibodies, Biopharmaceuticals and Edible Vaccines HENRY DANIELL, OLGA CARMONA-SANCHEZ and BRITTANY E. BURNS	113
8.1	Introduction	113
8.2	Expression of Therapeutic and Human Proteins in Plants	114
8.3	The Transgenic Chloroplast System	114
8.3.1	Chloroplast-derived Human Antibodies	116
8.3.2	Chloroplast-derived Biopharmaceuticals	118
8.3.2.1	Human Serum Albumin	118
8.3.2.2	Human Insulin-like Growth Factor-1	119
8.3.2.3	Human Interferon (IFN2b)	119
8.3.2.4	Anti-Microbial Peptides (AMPs): MSI-99	122
8.3.3	Chloroplast-derived Vaccine Antigens	123
8.3.3.1	Cholera Toxin B Subunit (CTB)	123
8.3.3.2	Bacillus anthracis Protective Antigen	124
8.3.3.3	Yersinia pestis F1~V Fusion Antigen	126
8.3.3.4	Canine Parvovirus (CPV) VP2 Protein	127
8.4	Advances in Purification Strategies for Biopharmaceuticals	129
8.5	Conclusion	131
	Acknowledgements	131
	References	131
9	Plant-derived vaccines: progress and constraints GURUATMA KHALSA, HUGH S. MASON, CHARLES J. ARNTZEN	135
9.1	Introduction	135
9.2	Strategies for Vaccine Production in Plants	138
9.3	The Biomanufacture of Vaccines	139
9.3.1	Advantages of Plants	139
9.3.2	Oral Delivery and Mucosal Immune Responses	140
9.3.4	Examples of Antigens Produced in Plants	140
9.3.5	Targeting Antigens to Specific Tissues	140
9.3.6	Expression Systems	141
9.3.7	Mucosally-targeted Fusion Proteins	142
9.3.8	Forming Multivalent and Multicomponent Vaccines	143
9.3.9	Stability and Processing	151
9.4	Clinical Trials with Plant-derived Vaccines	151
9.4.1	Enterotoxic E. coli and Vibrio cholerae	152
9.4.2	Norwalk Virus	152
9.4.3	Hepatitis B Virus	153
9.4.4	Rabies Virus	153
9.5	Issues and Challenges	154
9.5.1	Development and Licensing of Plant-derived Vaccines	154
9.5.2	Confronting GM Food Issues	154
	References	155
10	Production of Secretory IgA in Transgenic Plants DANIEL CHARGELEGUE, PASCAL M.W. DRAKE, PATRICIA OBREGON and JULIAN K.-C. MA	159
10.1	Introduction	159
10.2	Antibodies	159
10.2.1	Mucosal Antibodies	160
10.2.2	Structure and `Natural’ Production of SIgA	160
10.2.3	Passive Immunization with SIgA	162
10.2.4	Production of Recombinant SIgA	162
10.3	Production of Recombinant SIgA in Plants	163
10.3.1	Production of Full-length Antibodies in Plants	163
10.3.2	Production of Multimeric Antibodies: SIgA	165
10.3.3	Glycosylation of Antibodies in Transgenic Plants	166
10.3.4	Plant Hosts	167
10.4	Conclusions	167
	References	168
11	Production of Spider Silk Proteins in Transgenic Tobacco and Potato JÜRGEN SCHELLER and UDO CONRAD	171
11.1	Introduction	171
11.1.1	Structure and Properties of Spider Silk	171
11.1.2	Strategies for the Production of Recombinant Spider Silk Proteins	173
11.1.3	Applications of Spider Silk Proteins	174
11.1.3.1	Synthetic Spider Silk Fibers: `Natural’ vs Artificial Spinning Strategies	174
11.1.3.2	Synthetic Spider Silk Proteins for the In Vitro Proliferation of Anchorage-dependent Cells	175
11.1.4	Molecular Farming: Plants as Biofactories for the Production of Recombinant Proteins	175
11.2	Spider Silk and Spider Silk-ELP Fusion Proteins from Plants: Expression, Purification and Applications	176
11.2.1	Spider Silk-ELP Expression in Transgenic Tobacco and Potato	176
11.2.2	Purification of Spider Silk-Elastin Fusion Proteins by Heat Treatment and Inverse Transition Cycling	177
11.2.3	Applications of Spider Silk-ELP Fusion Proteins in Mammalian Cell Culture	178
11.3	Discussion	179
	References	180
12	Gene Farming in Pea Under Field Conditions MARTIN GIERSBERG, ISOLDE SAALBACH and HELMUT BÄUMLEIN	183
12.1	Introduction	183
12.2	Procedures for Foreign Protein Expression in Transgenic Pea Seeds	184
12.2.1	Plant Material, Transformation and Field Growth	184
12.2.2	Transformation Vectors and Analysis of Transgenic Plants	185
12.3	Expression of ?-Amylase in Transgenic Pea Seeds	185
12.4	Conclusions	188
12.5	Acknowledgements	189
	References	190
13	Host Plants, Systems and Expression Strategies for Molecular Farming RICHARD M. TWYMAN	191
13.1	Introduction	191
13.2	Host Species for Molecular Farming	194
13.2.1	Leafy Crops	194
13.2.1.1	Tobacco (Nicotiana tabacum)	194
13.2.1.2	Tobacco (Nicotiana benthamiana)	195
13.2.1.3	Alfalfa (Medicago sativa)	195
13.2.1.4	White clover (Trifolium repens)	195
13.2.1.5	Lettuce (Lactuca sativa)	196
13.2.1.6	Spinach (Spinacia oleracea)	196
13.2.1.7	Lupin (Lupinus spp.)	196
13.2.2	Dry Seed Crops	196
13.2.3	Fruit and vegetable crops	199
13.2.4	Oilcrops	201
13.2.5	Unicellular Plants and Aquatic Plants Maintained in Bioreactors	203
13.2.6	Non-cultivated Model Plants	204
13.3	Expression systems for molecular farming	205
13.3.1	Transgenic plants	206
13.3.2	Transplastomic plants	207
13.3.3	Virus-infected plants	207
13.3.4	Transiently transformed leaves	208
13.3.5	Hydroponic cultures	209
13.3.6	Hairy roots	209
13.3.7	Shooty teratomas	210
13.3.8	Suspension cell cultures	210
13.4	Expression strategies and protein yields	210
13.5	Conclusions	212
	References	213
14	Downstream Processing of Plant-derived Recombinant Therapeutic Proteins JUERGEN DROSSARD	217
14.1	Introduction	217
14.2	Similarities and Differences in the Processing of Pharmaceutical Proteins from Different Sources	218
14.3	Process Scale	220
14.4	The Individual Steps of a Downstream Process	221
14.4.1	Initial Processing and Extraction	222
14.4.2	Chromatographic Purification	224
14.5	Regulatory Requirements for Downstream Processing of Plant-derived Pharmaceutical Products	228
	References	230
15	Glycosylation of Plant-made Pharmaceuticals VÉRONIQUE GOMORD, ANNE-CATHERINE FITCHETTE, PATRICE LEROUGE and LOÏC FAYE	233
15.1	Introduction	233
15.2	Plant Cells can Reproduce the Complexity of Mammalian Proteins	233
15.3	Plant-made Pharmaceuticals and their Native Mammalian Counterparts Contain Structurally-distinct N-linked Glycans	238
15.4	Plant-made Pharmaceuticals Possess Immunogenic N-glycans	241
15.5	Current Strategies to Eliminate Immunogenic N-glycans from Plant-made Pharmaceuticals	242
15.6	Towards Humanized N-glycans on PMPs Through the Expression of Mammalian Glycosyltransferases in the Plant Golgi Apparatus	245
15.7	Concluding Remarks	248
15.8	Acknowledgements	248
	References	248
16	Biosafety Aspects of Molecular Farming in Plants ULRICH COMMANDEUR and RICHARD M. TWYMAN	251
16.1	Introduction	251
16.2	Transgene Spread	252
16.2.1	Classes of Foreign DNA Sequences in Transgenic Plants	252
16.2.2	Mechanisms of Transgene Pollution – Vertical Gene Transfer	253
16.2.3	Mechanisms of Transgene Pollution – Horizontal Gene Transfer	253
16.3	Combating the Vertical Spread of Transgenes	254
16.3.1	Choosing an Appropriate Host	254
16.3.2	Using Only Essential Genetic Information	255
16.3.3	Elimination of Markers After Transformation	257
16.3.4	Containment of Essential Transgenes	259
16.4	Unintended Exposure to Recombinant Proteins	261
16.4.1	Environmental Risks of Unintended Exposure	261
16.4.2	Addressing the Risks of Unintended Exposure	262
16.4.2.1	Controlling Transgene Expression	262
16.4.2.2	Controlling Protein Accumulation and Activity	263
16.4.2.3	Contamination of the Food Chain During Processing	263
16.5	Conclusions	264
	References	265
17	A Top-down View of Molecular Farming from the Pharmaceutical Industry: Requirements and Expectations FRIEDRICH BISCHOFF	267
17.1	Introduction	267
17.2	Industrial Production: The Current Situation	267
17.3	Expectations	270
17.4	Requirements	273
17.4.1	Equivalence of the Recombinant Product to the Original Protein	273
17.4.2	Processing in the Endoplasmic Reticulum (ER)	274
17.4.3	Glycosylation in the Golgi	275
17.4.4	Differential Glycosylation – Implications on Immunogenicity of vaccines	277
17.4.5	Glycosylation and Stability	277
17.4.6	Equivalence of Enzymes	279
17.4.7	Degradation	279
17.4.8	Efficacy in Clinical Trials	280
17.4.9	The Optimal Production System	283
17.4.10	Post-harvest expression	285
17.4.11	Purification	285
17.5	Conclusions	287
	References	287
18	The Role of Science and Discourse in the Application of the Precautionary Approach KLAUS AMMANN	291
18.1	Introduction	291
18.2	Other Roots to Problems with the Precautionary Approach	292
18.2.1	The Roots of the Precautionary Approach and Environmental Debate	292
18.2.2	Discussion About the PA is Too Closely Related to Factual Knowledge Alone	294
18.3	The First and Second Generation Systems Approaches	295
18.3.1	First Generation Systems Approach	295
18.3.2	Second Generation Systems Approach	295
18.4	How to Solve Wicked Problems in Biotechnology and the Environment	298
18.5	How to Achieve Such Demanding Planning Goals	299
18.6	There is no Scientific Planning	299
18.7	Outlook	300
	Bibliography	301

	Subject Index	303

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