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The appropriate viral vector for a given gene therapy application depends on the duration of gene expression desired, the cell type to be transduced, the immune pro-tected or unpropro-te

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Advances in molecular biology, cell

biology, and polymer chemistry are

providing novel approaches for

treating musculoskeletal disorders

These advances are facilitating

bio-logic approaches that complement

and could even replace traditional

biomechanical techniques in

ortho-paedic surgery The explosion of

research has resulted in a myriad

of potential applications for these

innovative biologic approaches

Many probably will never reach

clinical fruition; however, effective

implementation of only a few may

revolutionize the way we manage

certain musculoskeletal disorders

Clearly, orthopaedic surgeons must

have a fundamental understanding

of the terminology (Table 1) and

techniques of these biologic

ad-vances, and of current research, to

be able to interpret the literature,

direct future investigations, and

best serve their patients

Tissue Engineering

Tissue engineering is the science of creating living tissue to replace, repair, or augment diseased tissue.1 Tissue engineering thus refers to a broad variety of techniques The engineered tissue is created either entirely in vivo or, in an ex vivo technique, is created in vitro and subsequently implanted into the patient Regardless of the tech-nique, however, tissue engineering requires three components: a growth-inducing stimulus (growth factor), responsive cells, and a scaffold to support tissue formation

Growth Factors and Gene Therapy

Cytokines are small, soluble pro-teins that influence cell behavior

A subset of cytokines, known as growth factors, promotes cell mito-sis The term growth factor,

how-ever, often is used generically in reference to any number of cyto-kines that promote cell division, maturation, and/or differentiation Recent advances in molecular biology have facilitated the genetic sequencing of various cytokines Consequently, the genetic sequences can be inserted into cells of labora-tory animals, producing large quan-tities of “recombinant” human cyto-kines A particular cytokine often has multiple, diverse effects, and these can vary depending on the different cell types

Determining the influence and pharmacokinetics of various cyto-kines is an area of active, ongoing investigation In vitro research typ-ically is performed to identify the optimal cytokine for a particular

Dr Musgrave is Chief Resident, Department of Orthopaedic Surgery, University of Pittsburgh Medical Center, Pittsburgh, Pa Dr Fu is David Silver Professor and Chairman, Department of Orthopaedic Surgery, University of Pittsburgh Medical Center, Pittsburgh Dr Huard is Associate Professor, Departments of Orthopaedic Surgery, Molecular Genetics and Biochemistry, and Bioengineering, University of Pittsburgh, Pittsburgh.

Reprint requests: Dr Huard, Room 4151, Rangos Research Center, 3705 Fifth Avenue, Pittsburgh, PA 15213-2583.

Copyright 2002 by the American Academy of Orthopaedic Surgeons.

Abstract

A new biologic era of orthopaedic surgery has been initiated by basic scientific

advances that have resulted in the development of gene therapy and tissue

engi-neering approaches for treating musculoskeletal disorders The terminology,

fundamental concepts, and current research in this burgeoning field must be

understood by practicing orthopaedic surgeons Different gene therapy

approaches, multiple gene vectors, a multitude of cytokines, a growing list of

potential scaffolds, and putative stem cells are being studied Gene therapy and

tissue engineering applications for bone healing, articular disorders,

interverte-bral disk pathology, and skeletal muscle injuries are being explored Innovative

methodologies that ensure patient safety can potentially lead to many new

treat-ment strategies for musculoskeletal conditions.

J Am Acad Orthop Surg 2002;10:6-15

Gene Therapy and Tissue Engineering

in Orthopaedic Surgery

Douglas S Musgrave, MD, Freddie H Fu, MD, and Johnny Huard, PhD

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application The optimal method

by which to deliver the cytokine in

vivo also must be determined The

simplest method is to inject or

im-plant the recombinant protein

di-rectly into the anatomic region of

interest However, the inherently

short half-lives of recombinant

pro-teins and the dilutive effects of

body fluids limit this technique

Consequently, the injected

cyto-kines may not be present in

ade-quate concentrations and at the

critical time periods to effect a

maximal response Therefore,

in-vestigators have devised two basic approaches to deliver temporary, sustained quantities of a desired protein

The first method is to impreg-nate the cytokine onto a polymer scaffold Such a method requires a thorough understanding of the chemical interactions between the protein and the scaffold, the bind-ing characteristics of the protein to the scaffold, and the pharmacoki-netics of both the protein and the scaffold The second method is to deliver the gene encoding for the

protein into the patient, resulting in

in vivo cytokine production When the gene is delivered only to a spe-cific region of the body and not systemically, the method is termed regional gene therapy

Gene therapy is simply the transfer of a particular gene into a cell so that the cell transcribes the gene into messenger ribonucleic acid (mRNA); the cell’s ribosomes then translate the mRNA into a protein (cytokine) To accomplish this, the particular gene must be packaged and inserted into the

Table 1

Common Terms Used in Gene Therapy

Term Definition

cDNA Complementary DNA DNA created from a protein’s mRNA by enzyme reverse transcriptase

Contains only exons

mRNA RNA that is produced by transcription of DNA This genetic material leaves the nucleus, moves to the

cytoplasm, and is translated by ribosomes into a protein

RNA A string of ribonucleotides that is similar in structure to DNA There are several classes of RNA,

including messenger RNA (mRNA), transfer RNA (tRNA), and ribosomal RNA (rRNA)

Nucleotide Building block of DNA and RNA comprising a base, a deoxyribose or ribose sugar, and a phosphate Exon A piece of DNA encoding a gene that is transcribed into mRNA and then translated into a protein

(that is, a nucleotide sequence of a gene that encodes for a specific protein)

Intron A piece of DNA of unknown function that is interspersed between exons (that is, a noncoding

nucleotide sequence of a gene)

Transcription The process of conversion of DNA into RNA (that is, creation of an mRNA from DNA)

Transduction Gene transfer using viral vectors

Transfection Gene transfer using nonviral vectors

Translation The process of conversion of mRNA into an amino acid sequence (that is, creation of a protein from

mRNA by ribosomes)

Plasmid A self-replicating piece of DNA naturally found in bacteria and yeast Plasmids are commonly used to

carry foreign genes into cells for the production of recombinant DNA pharmaceuticals and in gene expression studies

Liposome A gene-delivery system based on the production of lipid bodies that contain pieces of DNA These

lipid-DNA complexes fuse with the cell surface and are internalized by phagocytosis, depositing the encapsulated DNA into the target cells

Promoter A regulatory element in DNA that functions to induce transcription of a gene

Cytokines Small proteins that influence cell behavior

Ex vivo/In vitro

gene transfer Genetic manipulation of cells outside the host (body)

In vivo gene

transfer Genetic manipulation of cells within the host (body)

Vector A disabled virus or DNA structure used as a vehicle to transfer genes into cells

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is reverse-transcribed in the

labora-tory by the enzyme reverse

trans-criptase to create a complementary

deoxyribonucleic acid (cDNA)

Because the cDNA is created

di-rectly from mRNA, the noncoding

nucleotide sequences (introns) of

the gene are not present, leaving

only the protein-coding nucleotide

sequences (exons) This fact

distin-guishes a protein’s cDNA from its

DNA sequence found naturally,

which includes both introns and

exons

The cDNA is inserted into a

cir-cular plasmid with various non–

amino acid-coding sequences at its

leading and trailing ends (Fig 1, A)

A plasmid is a self-replicating,

cir-cular piece of DNA that is used to

transfer specific genes into cells

The cDNA plasmid is then digested

with restriction enzymes and

in-serted into a larger plasmid

contain-ing a promoter sequence (among

other sequences), thereby creating

an expression plasmid (Fig 1, B)

The promoter sequence induces

transcription of the cDNA once

gene transfer into a cell has

oc-curred Frequently, the promoter is

constitutive, meaning it is always

“on” and the gene will be

tran-scribed continuously until the gene

is lost The cytomegalovirus

pro-moter is a commonly used

consti-tutive promoter Investigators are

currently developing regulated

pro-moters, which can be turned “on”

or “off” with an exogenous stimulus

(i.e., a drug) Regulated promoters

are obviously desirable so that gene

expression may occur only at the

desired, critical time periods Once

a functional expression plasmid

has been created, one must

deter-mine the optimal vector with which

to transport the expression plasmid

safely and efficiently into the

de-sired cells

Vectors are vehicles that

facili-tate transfer of a particular gene

into cells Vectors are subdivided

into viral and nonviral vectors The most commonly used nonviral vec-tors are liposomes Liposomes are phospholipid vesicles capable of fusing with a cell membrane,

there-by delivering their contents (ex-pression plasmid) into the cell

Liposomes are relatively cheap and nonpathogenic, but their use results

in inferior gene-transfer efficiency compared with viral vectors Other nonviral methods include the gene gun (DNA loaded onto gold beads injected into the cell using a helium

“gun”), DNA conjugates (DNA conjugated to certain polycations that improve DNA adherence to cell membranes, thereby improving gene transfer), gene-activated matrices (matrices, or scaffolds, im-pregnated with expression plas-mids), and nonviral-viral hybrids (viral DNA sequences that facilitate longer gene expression or integra-tion into host cell DNA spliced into

an expression plasmid)

Successful gene transfer using nonviral vectors is termed trans-fection Alternatively, successful

gene transfer using viral vectors is termed transduction Using current methods, the efficiency of nonviral transfection is generally inferior to that of viral-mediated transduction Therefore, despite the nonpatho-genic nature and economic advan-tages of nonviral vectors, most cur-rent gene therapy applications use viral vectors

Viruses are highly efficient at infecting cells to deliver genetic material Various viruses have dif-ferent advantages and disadvan-tages when applied to gene therapy (Table 2) The most commonly used viruses in current gene therapy ap-plications include adenoviruses, retroviruses, herpes simplex viruses (HSVs), and adeno-associated viruses (AAVs) Certain viral DNA se-quences are often removed to ren-der the viral replication defective before gene therapy applications Adenoviruses have the advan-tage of infecting both dividing and some nondividing cells, thereby achieving a high level of transient gene expression The adenovirus

G E

N E

restriction sites

polyadenylation

restriction sites

restriction sites

cDNA

restriction sites

A A

G E NE

P R O M

OT ER

A M

P

N E O

EXPRESSION PLASMID

Figure 1 A, Complementary DNA (cDNA) consisting of desired gene and restriction enzyme sites in circular form B, An expression plasmid is created by inserting the desired

gene into plasmid containing other sequences A promoter initiates transcription of the gene, and the polyadenylation sequence helps stabilize the mRNA Ampicillin (AMP) and neomycin (NEO) resistance genes also may be present to allow in vitro selection of trans-duced or transfected cells.

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genome exists as an episome within

the cell’s nucleus, not being

inte-grated into the cell’s genome

Low-level production of adenoviral

anti-gens often results in an immune

response to infected cells, resulting

in loss of gene expression after

sev-eral weeks Research is ongoing to

develop “gutted” adenoviral

vec-tors, whereby the majority of viral

protein expression is eliminated to

minimize the immune response to

adenoviral antigens

Retroviruses are small RNA

viruses that, once inside the cell,

have their RNA transcribed by the

enzyme reverse transcriptase into

stranded DNA The

double-stranded DNA enters the cell

nu-cleus and is integrated, although at a

random site, into the cell’s genome

Consequently, the therapeutic gene

is expressed for the life of the cell

The disadvantages of retroviral

vec-tors are that they infect and

trans-duce only actively dividing cells

and that their DNA is randomly

integrated into the host cell genome

This random integration

theoreti-cally can alter oncogene expression

and adversely affect cell behavior

HSV is capable of infecting both

dividing and nondividing cells,

car-rying large amounts of DNA,

infect-ing many different cell types, and establishing latency in neuronal cells First-generation HSV vectors demonstrated toxicity in certain cell types and expressed significant anti-genicity, resulting in only transient gene transfer in certain cell types

Next-generation HSV vectors thus are being engineered to minimize these drawbacks

AAV has the advantages of infecting nondividing cells and of stable integration of its DNA into a specific site on chromosome 19, which appears to be nonpatho-genic.2 Furthermore, AAV is not known to cause any disease.3 The main disadvantage of AAV is its capacity to accommodate only small amounts of exogenous DNA, on the order of 5.2 kilobase (kb) pairs

The appropriate viral vector for

a given gene therapy application depends on the duration of gene expression desired, the cell type to

be transduced, the immune pro-tected or unpropro-tected environment

in which the cells reside, and the method chosen to ensure patient safety (i.e., in vitro and/or in vivo safety testing)

As mentioned, two fundamental approaches to gene therapy exist: in vivo (direct) and ex vivo (outside

the host) (Fig 2) The in vivo ap-proach consists of directly injecting

or implanting the gene-carrying vector into the patient This ap-proach is attractive for its technical simplicity although it is limited by the inability to perform in vitro safety testing on the transduced cells The alternative ex vivo approach consists of isolating cells in vitro from a tissue biopsy taken from the patient The cells are expanded and then transfected or transduced in the laboratory The genetically altered cells can be tested in vitro for both successful gene transfer and abnormal behavior before they are reimplanted in the patient After in vitro testing, the cells are introduced into the patient so that they may produce the genetically specified protein The ability to test for successful gene transfer and abnormal cellular behavior before cell implantation is an advantage of the ex vivo approach However, the

ex vivo method is technically more complex than the in vivo approach

Mesenchymal Stem Cells

Growth factors are capable of acting on many different cell types However, mesenchymal stem cells (MSCs) are the ideal cell type for

Table 2

Common Viral Vectors

Adenovirus Infects dividing and nondividing cells Viral antigens can induce immune response

Infects many different cell types DNA remains episomal; does not integrate Straightforward vector production

Retrovirus DNA integrates into cell’s genome Random DNA integration

Accomodates 8 kb Infects only dividing cells Straightforward vector production

Herpes simplex virus Infects dividing and nondividing cells Toxicity

Accomodates >35 kb Transient expression in certain cell types Adeno-associated virus Infects nondividing cells Accomodates only 5.2 kb

Nonpathogenic Difficult vector production Infects many different cell types

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therapy approaches MSCs are

rest-ing stem cells capable of

differenti-ation into multiple connective

tis-sue lineages (such as muscle, bone,

ligament, tendon, and cartilage).4,5

MSCs are not to be confused with

hematopoietic stem cells, which are

obtained from fetal umbilical cord

blood and are capable of

differenti-ation into multiple hematopoietic

lineages (such as megakaryocytes,

erythrocytes, lymphocytes).6

The existence of MSCs facilitates

seemingly endless tissue engineering

possibilities Theoretically, an MSC

could be stimulated to undergo

dif-ferentiation down a desired lineage,

thereby recreating a certain tissue for

therapeutic use Isolation of fully

differentiated cells (i.e.,

chondro-cytes) for a certain tissue engineering

application can be hindered by

donor site morbidity and limited cell

availability The use of MSCs would

obviate the need to obtain fully

dif-ferentiated cells for a specific tissue

engineering application

MSCs are thought to reside in the

bone marrow and thus are obtained

by bone marrow biopsy Resting

osteoprogenitor stem cells reside in

the bone marrow stroma, and bone

marrow–derived MSCs have been

used in animal studies to heal

carti-lage defects7as well as produce

cel-lular elements that could modify the

clinical course of muscle diseases.8

Additionally, the feasibility of using

MSCs in gene therapy has been

de-monstrated in animal models.9

Furthermore, the effective human

clinical use of MSCs to treat

neo-nates with osteogenesis imperfecta

has been reported.10,11 Clinical

im-provement was demonstrated in

neonates with osteogenesis

imper-fecta who received allogeneic bone

marrow transplantation aimed at

replacing their abnormal MSCs

MSCs also can reside outside the

bone marrow: evidence supports

the existence of resting stem cells in

skeletal muscle that are capable of

either myogenic or osteogenic dif-ferentiation.12-14 The availability of skeletal muscle and the relative ease

of cell isolation make skeletal mus-cle an attractive source of potential stem cells The relationship of these muscle-derived stem cells and MSCs still is unclear and remains to be in-vestigated fully

In addition, stem cells possibly may reside in other tissues, such as skin, brain, kidney, or perivascular tissue Further research must be conducted to elucidate fully the existence and residence of stem cells and to refine efficient isolation tech-niques

Scaffolds

A scaffold to support tissue growth is the final component of any tissue engineering approach

This scaffold can take the form of

in situ host tissue (e.g., meniscus), transplanted host tissue (e.g., skele-tal muscle flap), naturally derived polymers (such as collagen and hyaluronic acid), synthetic

poly-mers (such as poly[L-lactic acid] [PLLA], polyglycolic acid [PGA], and poly[DL-lactic-co-glycolic acid] [PLGA]), or injectable polymers that cross-link in situ (such as algi-nate and polyethylene oxide [PEO]) (Table 3) Scaffolds influence cell recruitment, cell containment, cell adherence, diffusion of nutrients to the cells, delivery of growth fac-tors, and cell behavior An ideal scaffold provides for uniform dis-tribution of cells throughout its three-dimensional lattice, facilitates efficient diffusion of biochemical molecules, and undergoes degrada-tion at the same rate that it can be replaced by host tissue The latter characteristic dictates the species-specific suitability of a scaffold Certain scaffolds can be impreg-nated with expression vectors,

there-by creating gene-activated matrices (GAMs) In addition to fulfilling the previously mentioned scaffold char-acteristics, GAMs can provide for nonviral direct gene transfer, thereby avoiding the immunologic risks of

Figure 2 Two basic approaches of gene therapy exist Ex vivo gene therapy consists of

cell harvest and expansion in culture, in vitro transduction or transfection of the isolated cells using the appropriate vectors, and reimplantation of the transduced cells into the patient In vivo gene therapy consists of direct injection of the vectors into the patient.

Cell harvest and culture

In vitro transduction

Virus

Reimplantation

of transduced cells

Direct injection

of vector

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viral vectors Host-tissue scaffolds

are attractive because they obviate

the need for a foreign body, may

contain responsive cells, and

poten-tially can be vascularized However,

donor-site morbidity and lack of

available tissue are prohibitive

fac-tors Naturally derived polymers are

attractive because they may be

remodeled by host cells to provide

space for growing tissue Collagen

menisci constructed from bovine

Achilles tendon type I collagen were

implanted into eight patients, with

encouraging results at second-look

arthroscopy and 2-year clinical

follow-up.15 The drawbacks of

natu-rally derived polymers are ensuring

pathogen removal and potential

lim-ited supply

Synthetic polymers, in contrast,

can be mass-produced and

de-signed specifically for a particular

application Their molecular

com-position can be modified to affect

scaffold-cell interactions and

scaf-fold degradation The presence of a

foreign body, however, may result

in an inflammatory reaction, to the

detriment of tissue growth, and

increase the risk of infection

Syn-thetic polymers have been used to

create composite phalanges by

seeding bovine chondrocytes and

tenocytes onto sheets of PGA and

suturing these structures to bovine

periosteum wrapped around cova-lently linked PLLA and PGA.16 The composite structures were subcuta-neously implanted into athymic mice to facilitate growth No gene transfer was used After 20 weeks, the composite tissue resembled the gross and histologic appearance of human phalanges

The science of polymer chemistry governing the porosity, cell- and growth factor–binding characteris-tics, and degradation kinematics of synthetic polymers is beyond the scope of this article It must be noted, however, that, given each scaffold’s inherent advantages and disadvan-tages, the scaffold for a given tissue engineering application must be cho-sen carefully

Specific Tissues Bone

The existence of bone-inducing growth factors has long been rec-ognized.17 These bone-inducing growth factors, termed bone mor-phogenetic proteins (BMPs), are members of the transforming growth factor-β(TGF-β) superfamily Recom-binant human BMP-2 (rhBMP-2)18 has been implanted directly on var-ious carriers and can heal critically sized bone defects in animal

mod-els.19 Likewise, other BMPs, such

as BMP-320and BMP-7 (osteogenic protein-1),21have proved to be effica-cious in animal models To achieve sustained BMP delivery, investiga-tors have developed both in vivo and ex vivo gene therapy approaches using BMPs Direct adenovirus-mediated approaches delivering BMP-222,23 and BMP-9,24 as well as

a direct GAM-mediated approach delivering BMP-4,25 have been reported Ex vivo approaches to deliver BMP-2 have used rodent bone marrow stromal cell lines,26,27 primary rodent bone marrow stro-mal cells,28,29primary rodent skele-tal muscle–derived cells,12,29primary human skeletal muscle–derived cells,30primary rabbit articular chon-drocytes,29 primary rabbit perios-teal cells (BMP-7),31 and primary rabbit skin fibroblasts.29 For clin-ical applications, primary autolo-gous cells (cells isolated from the host) are preferable to cell culture lines, which are allogeneic or xeno-geneic and may have tumorigenic and immunogenic potential The direct, in vivo gene therapy approach using BMPs has been ap-plied in rodent spine fusion mod-els,22,24 whereas the ex vivo ap-proach has been applied to healing critically sized rodent bone de-fects.13,26,28 In the spine fusion

Table 3

Categories of Scaffolds

Host tissue Responsive cells Donor site morbidity

(e.g., meniscus, skeletal muscle flap) Viability and potential vascularity Limitation on amount of tissue

No foreign body Naturally derived polymers Remodeling potential Pathogen removal must be assured (e.g., collagen, hyaluronic acid) No donor site morbidity Potential limited supply

Synthetic polymers Mass production Foreign body (infection risk)

(e.g., PLLA, PGA, PLGA) Custom designed Possible inflammatory reaction Injectable synthetic polymers Mass production Foreign body (infection risk)

(e.g., alginate, PEO) Can fill complex spaces In vivo chemical reaction

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directly injected into the paraspinal

musculature,22,24resulting in

para-spinal bone formation present at 3

weeks after injection In the ex vivo

approaches, the transduced cells

were seeded onto collagen sponges13

or demineralized bone matrix26,28as

scaffolds, resulting in improved

radiographic healing of both

calvar-ial13and long bone defects.26,28

A novel osteoinductive protein

termed LIM mineralization

protein-1 (LMP-protein-1) has been used in an ex

vivo gene therapy model for spine

fusions.32 Because LMP-1 is an

intracellular signaling molecule

(regulated by BMP-6), the use of

LMP-1 requires gene transfer Bone

marrow stromal cells transfected

with LMP-1, seeded onto a scaffold

of nonosteogenic devitalized bone

matrix, and implanted into the

paraspinal area of rats resulted in a

100% rate of spine fusion compared

with no fusion in the controls.32

Which cell type to use in ex vivo

gene transfer of osteogenic proteins

remains unresolved The ideal cell

type is expendable, easily isolated

from the patient, and amenable to

efficient transduction and protein

secretion; possesses

osteocompe-tence17; and has favorable survival

characteristics when reimplanted

into the patient Bone marrow

stro-mal cells26,28,29 and skeletal

mus-cle–derived cells12-14,29both possess

osteocompetence and have been

used successfully in ex vivo

osteo-genic protein gene transfer

Fi-nally, as mentioned previously, the

development of composite

pha-langes may be possible without

gene therapy using a combination

of cell transplantation and polymer

scaffolds.16

Growth plate disorders are

an-other possible skeletal gene therapy

application In a rabbit physeal

injury model, direct

adenovirus-mediated gene transfer of

insulin-like growth factor-1 (IGF-1) was

shown to inhibit subsequent

phy-direct BMP-2 gene transfer promoted premature physeal closure and sub-sequent growth disturbance.33 These results warrant further investigation into the use of IGF-1 gene transfer to inhibit physeal closure after physeal fractures and into the development

of physiodesis using BMP-2 gene transfer

Articular Structures

Patients with a variety of articu-lar disorders are potential candi-dates for gene therapy and tissue engineering applications Innova-tive approaches for treating arthri-tis, chondral and osteochondral de-fects, meniscal tears, and ligament injuries currently are being investi-gated

Arthritis

Arthritis was the first nonlethal disease for which a human gene therapy trial was approved and undertaken.34 This trial consisted of the ex vivo transfer of the inter-leukin-1 (IL-1) receptor antagonist protein (IRAP or IL-1RA) to syno-viocytes obtained from six post-menopausal women with severe rheumatoid arthritis A retroviral vector was used to transduce the synoviocytes with IRAP, a protein that inhibits the arthritogenic cy-tokine IL-1 The transduced syn-oviocytes were injected into the patients’ metacarpophalangeal (MCP) joints 1 week before sched-uled MCP joint arthroplasty At the time of MCP joint arthroplasty, the joints, synovium, and synovial fluid were harvested for analysis The trial was designed to establish feasi-bility and safety but not necessarily efficacy No adverse reactions in the patients have been reported to date, and lifelong clinical follow-up continues

Animal studies have established that direct adenoviral-mediated gene transfer of a soluble IL-1 recep-tor and a soluble tumor necrosis

matrix degradation in rabbit knees with antigen-induced arthritis.35 Combining multiple antiarthritic proteins in gene therapy applica-tions may lead to additive or syner-gistic effects

Cartilage

Different approaches have been applied to chondral and osteochon-dral defects RhBMP-2 regulates the behavior of costochondral growth plate chondrocytes36and maintains the phenotype of articular chon-drocytes in cell culture.37 Collagen sponges impregnated with rhBMP-2 improve the healing of rabbit full-thickness cartilage defects.38 The feasibility of using articular chon-drocytes in ex vivo BMP-2 gene transfer has been established.16 The effects of in vitro gene transfer of IGF-1, BMP-2, and TGF-β to rabbit articular chondrocytes have recently been investigated.39 Gene transfer

of BMP-2 was a potent stimulus for proteoglycan synthesis in the pres-ence of IL-1 Gene transfer of IGF-1 was a strong stimulus for collagen and noncollagenous protein synthe-sis Other factors, such as BMP-7, cartilage growth and differentiation factors, and various transcription factors, also may play a role carti-lage healing.40

Cell transplantation of autolo-gous chondrocytes injected into chondral defects and covered with

a periosteal flap has shown some promise clinically, especially for femoral condyle lesions.41 Cell transplantation using various natu-rally derived scaffolds, synthetic scaffolds, injectable scaffolds, and autologous scaffolds42 is being in-vestigated in animal models.43

In addition to other tissues, bio-reactor vessels may hold promise for tissue engineered cartilage Bioreactor vessels are cell culture containers specially designed to facilitate manipulation of the cells’ environment Environmental

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fac-tors, such as culture medium flow

and mechanical stress, can be

ma-nipulated to influence cell behavior

and tissue growth Bioreactor

ves-sels can be used both to engineer

cartilage for possible implantation

and provide a controlled in vitro

environment for the study of

chon-drogenesis.44 Size limitations of

bioreactor vessels are the current

restraints to clinical application

The optimal combination of growth

factor, delivery method, cells, and

scaffold to heal cartilage injuries is

the subject of ongoing investigation

Meniscus

The possibility of creating in

vitro a custom replacement

menis-cus using scaffolds,15 cells, and/or

gene therapy for subsequent in vivo

implantation is intriguing

Alterna-tively, meniscal cells might be

mod-ulated in vivo using gene therapy to

promote healing of certain injuries

Meniscal cells are amenable to gene

transfer of both marker genes and

various growth factor genes using

either in vivo or ex vivo gene

ther-apy, with gene expression persisting

for up to 6 weeks.45-47 Successful ex

vivo gene transfer has been

accom-plished using either myoblasts46,47

or meniscal cells.45 Implanted

me-niscal scaffolds constructed of

bo-vine collagen appear to be replaced

by host tissue mimicking a human

meniscus.15 Research is ongoing

into the preferred growth factors to

promote meniscal healing,

tech-niques to improve long-term gene

expression, and the optimal scaffold

needed to create new menisci

Ligaments

Gene therapy techniques also are

being applied to ligaments

β-Galactosidase (lacZ) gene transfer to

ligament has proved to be feasible in

animal models by either the in vivo

or ex vivo approach, using both

ade-novirus as well as retrovirus Ex

vivo gene transfer to ligaments has

been successfully achieved using either ligament fibroblasts48,49 or skeletal muscle myoblasts.49 Many growth factors, such as basic fibro-blast growth factor, platelet-derived growth factor (PDGF), vascular endothelial growth factor, IGF-1 and -2, TGF-β, and BMP-12, may play roles in ligament healing Data sug-gest that PDGF stimulates cell divi-sion and migration, whereas TGF-β and the IGFs promote extracellular matrix synthesis.50 Direct gene transfer of PDGF-β using a viral-liposome conjugate vector into rat patellar ligament resulted in initial improved angiogenesis and subse-quent enhanced extracellular matrix synthesis.51 Studies to improve liga-ment-bone healing using BMP-12 and to engineer ligament grafts in vitro are ongoing

Intervertebral Disk

Intervertebral disk nucleus pul-posus cells reside in an immuno-privileged site, which makes them potentially attractive targets for gene therapy approaches Direct adenovirus-mediated gene transfer

of lacZ into rabbit nucleus pulposus

cells in an in vivo model resulted in persistent gene expression for at least 12 weeks.52 A similar approach has been used to transfer the human TGF-β1 gene to rabbit nucleus pul-posus cells in vivo.53 The cells were harvested 1 week later and in vitro assays were performed TGF-β1 gene transfer resulted in a 30-fold increase in active TGF-β1, a fivefold increase in total TGF-β1, and a 100%

increase in proteoglycan synthesis

The results suggest that gene ther-apy to treat degenerative disk dis-ease warrants further investigation

Skeletal Muscle

Both in vivo and ex vivo forms of gene therapy to skeletal muscle for various types of inherited diseases, such as Duchenne muscular dystro-phy, have been investigated for

many years.54 In clinical trials, autol-ogous myoblast transplantation for Duchenne muscular dystrophy has proved to be safe,55although myo-blast survival is often suboptimal for inherited muscle diseases.54 Additionally, ongoing research into novel growth factors that might facilitate gene therapy approaches

to expedite the healing of acquired muscle injury has many potentially far-reaching applications IGF-1, basic fibroblast growth factor, and nerve growth factor have been shown in rodent models to improve muscle healing and strength (fast-twitch or tetanic strength) after con-tusion, laceration, and strain.56-58 However, myoblast transplantation combined with ex vivo gene therapy

is an even more attractive approach that can be used to deliver appropri-ate growth factors and responsive cells for acquired, traumatic muscle injuries

Summary

The biologic era of orthopaedic surgery promises to change the care

of musculoskeletal disorders in this century Extensive laboratory in-vestigations and preliminary clinical investigations have established the feasibility and promise of gene ther-apy and tissue engineering Re-search must continue to further un-derstanding of cytokines, gene delivery vectors, MSCs, and scaf-folds Pertinent issues, such as the optimal growth factors or genes, the timing and control of growth factor delivery, the optimal target cell, and the effectiveness of various scaf-folds, must be addressed before widespread clinical use can occur Most important, early clinical trials must establish patient safety before clinical efficacy is sought Respon-sible and innovative investigation will lead orthopaedic surgery into this new biologic era

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