Journal of Virological Methods 188 (2013) 153–160

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Journal of Virological Methods

jou rnal homepage: www.elsevier.com/locate/jviromet

Simple diffusion-constrained immunoassay for p24 with the sensitivity

of nucleic acid amplification for detecting acute HIV infection

Lei Chang, Linan Song, David R. Fournier, Cheuk W. Kan, Purvish P. Patel, Evan P. Ferrell, Brian A. Pink,

Kaitlin A. Minnehan, David W. Hanlon, David C. Duffy, David H. Wilson

Quanterix Corp, 113 Hartwell Ave, Lexington, MA 02421, USA

a b s t r a c t

Article history: Nucleic acid amplification techniques have become the mainstay for ultimate sensitivity for detecting

Received 8 May 2012

low levels of virus, including human immunodeficiency virus (HIV). As a sophisticated technology with

Received in revised form 22 August 2012

relative expensive reagents and instrumentation, adoption of nucleic acid testing (NAT) can be cost

Accepted 29 August 2012

inhibited in settings in which access to extreme sensitivity could be clinically advantageous for detection

Available online 2 October 2012

of acute infection. A simple low cost digital immunoassay was developed for the p24 capsid protein of HIV

based on trapping enzyme-labeled immunocomplexes in high-density arrays of femtoliter microwells

Keywords:

and constraining the diffusion of the enzyme–substrate reaction. The digital immunoassay was evalu-

Digital ELISA

Immunoassay ated for analytical sensitivity for HIV capsid protein p24, and compared with commercially available NAT

methods and immunoassays for p24, including 4th-generation /antigen combo assays, for early

Single molecule array

p24 detection of HIV in infected individuals. The digital immunoassay was found to exhibit 2000–3000-fold

HIV greater analytical sensitivity than conventional immunoassays reactive for p24, and comparable sensitiv-

ity to NAT methods. Assaying serial samples from 10 HIV-infected individuals, the digital immunoassay

detected acute HIV infection as early as NAT methods, and 7–10 days earlier than conventional immunoas-

says. Comparison of assay results between the digital immunoassay and a quantitative NAT method

2

from HIV infected serum exhibited a linear correlation R > 0.99. The data indicate that by constraining

diffusion of the signal generation step of a simple sandwich immunoassay and enabling the digital count-

ing of immunocomplexes, dramatic improvements in sensitivity to virus can be obtained to match the

sensitivity of NAT at a fraction of the cost.

© 2012 Elsevier B.V. Open access under CC BY-NC-ND license.

1. Introduction sensitivity detection of acute HIV infection. NAT has significantly

shortened the window between initial infection and its detection

Since the development of the polymerase chain reaction in through amplification of viral nucleic acid rather than detecting

the 1980s (Mullis et al., 1986), amplification of specific nucleic the presence of antibody following seroconversion (Busch, 2007).

acid sequences for genetic identification has become an indis- Although NAT has become the mainstay for detection of viruses,

pensible tool in medical research and diagnostics, including the the technology is relatively complex and high cost (Westreich et al.,

detection and diagnosis of infectious disease. With its imple- 2008), and can be cost inhibitory or prohibitive in settings where

mentation for blood screening in the US in 1999 (Busch et al., access to high sensitivity could be clinically advantageous. These

2000), nucleic acid amplification testing (NAT) for detection of scenarios include blood donor screening in lower-resource settings

viral pathogens has helped safeguard blood for transfusion by and clinical screening in higher incidence areas, where a significant

providing the most sensitive, economically feasible detection of number of cases of acute infection can be missed by less sensi-

infected blood donations possible. For HIV detection, NAT meth- tive immunoassay tests. Rapid detection and reporting of acute HIV

ods in use for blood screening are capable of analytical sensitivities infection represents a key opportunity to control the march of the

of approximately 60 HIV RNA copies/mL (30 viruses/mL) with indi- disease because viral transmission is 10 times more likely during

vidual donors (Assal et al., 2009; Proceix ULTRIO Product Insert, the acute phase than in the chronic phase (Wawer et al., 2005).

2011). In the clinical setting, NAT is the gold standard for high Immunoassays, on the other hand, are simpler and lower

cost than NAT, and can be deployed in more diverse environ-

ments. Conveniently, nature provides its own amplification of

∗ protein molecules from each virus in the form of to the

Corresponding author.

E-mail address: [email protected] (D.H. Wilson). virus and the viral . Immunoassays to HIV antibodies are

0166-0934 © 2012 Elsevier B.V. Open access under CC BY-NC-ND license. http://dx.doi.org/10.1016/j.jviromet.2012.08.017

154 L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160

straightforward, but they require the immune response and are genetically engineered Saccharomyces cerevisiae. One amino acid

unable to detect acute infection when antibodies are not yet (met) was inserted at the N-terminus to enable expression.

present. 3rd generation antibody assays (Constantine et al., 1994)

detect the earliest stage of the immune response (immunoglobu- 2.2. Cultured virus

lin M), reducing the serological window to approximately 3 weeks

(Fiebig et al., 2003). Viral proteins are present at the start of infec- The cultured HIV-1 Group M, Subtype B virus was purchased

tion, and each HIV particle produces approximately 2000 copies from Seracare Life Sciences (Part number PN242B, lot 9899P-10,

9 ®

of p24 capsid protein (NIAID, 2011), which works in favor of 1.15 × 10 HIV-1 RNA/mL by Roche AMPLICOR HIV-1 MonitorTM

immunoassay detection of acute infection. 30 viruses/mL equates Assay, v. 1.5) and diluted in delipidated, defibrinated human

to approximately 60,000 p24 molecules per mL, or a concentration serum (Seracare PN: HS-210). This virus is harvested from the

of 3 fg/mL. Unfortunately, current, state-of-the-art conventional culture supernatant without additional purification; therefore the

immunoassays are not sufficiently sensitive to detect these low standard could contain free p24 in addition to virion-incorporated

7

numbers. Immunoassays for p24 (Kontio, 1991) and 4th genera- p24. The diluted virus, at 1 × 10 copies HIV-1 RNA/mL, was inactiv-

tion combo assays that combine reactivity for p24 and anti-HIV ated by the addition of Triton X-100 to 0.4% and incubation of the

antibodies (Weber et al., 1998) can only detect the presence sample for 1 h at 37 C (Ukkonen et al., 1988).

of circulating p24 antigen after it has elevated to greater than

11,000–70,000 fg/mL (Marcel et al., 2011). While this can reduce 2.3. Seroconversion panels

the detection window another 4 or 5 days relative to serology

tests, immunoassays for p24 remain thousands-fold less sensi- Sets of seroconversion serum samples from HIV infected indi-

tive for virus than HIV RNA detection, which reduces the window viduals were purchased from SeraCare (PRB956, PRB958, PRB967,

an additional week or more relative to antigen detection (Fiebig RB968, PRB969, PRB972) and from Zeptometrix (ZM-HIV9016, ZM-

et al., 2003). Thus conventional immunoassays remain blind to HIV9018, ZM-HIV9024, ZM-HIV9031).

most of the acute phase window that is detectable by NAT. A simple

immunoassay capable of the sensitivity of NAT for HIV could repre- 2.4. Digital immunoassay

sent another step forward in broader, more cost effective detection

of acute HIV infection, which would help further control the spread Digitization of immunoassay analyte detection using single

®

of the disease. molecule arrays (Simoa ) has been described (Rissin et al., 2010,

Thermodynamically, antigen–antibody interactions should 2011). In brief, it involves performing a standard paramagnetic

enable sufficient sensitivity for detection of fg/mL concentrations microbead-based enzyme-linked immunosorbent assay (ELISA),

of antigen with little amplification. The problem for unamplified singulation of individual microbeads in high-density arrays of 50 fL

methods has been acquiring sufficient signal of this interaction. microwells, and confinement within the microwells of fluorescent

Conventional immunoassays employing optical signal detection product generated by enzymes labeling the immunocomplexes.

(such as chemiluminescence) require millions to thousands of Preventing diffusion of the fluorescent product out of the wells per-

millions of signal molecule events before signal can be reliably mits rapid buildup of fluorescence that is readily visualized with a

discerned by a luminometer or fluorometer. This limitation arises CCD camera. Immunocomplexes are labeled with ␤-galactosidase

because the ensemble signal is diluted in a typical reaction vol- (␤G) during the ELISA, and 30 s of conversion of enzyme substrate

ume of hundreds of microliters required for measurement. When (resorufin ␤-d-galactopyranoside, RGP) into fluorescent product

the signal molecules are constrained from diffusing beyond a small (resorufin) by a single molecule of ␤G provides sufficient signal

local volume, then lower numbers of signal molecules would be suf- to differentiate wells containing beads with an enzyme label from

ficient for optical detection due to their local high concentration. wells containing beads without an enzyme. At very low p24 con-

With an isolated enzyme-labeled antigen, the smaller the volume of centrations, Poisson statistics predict that bead-containing wells

confinement, the higher the local concentration of signal molecules in the array will contain either a single labeled p24 molecule or no

for detection. By confining enzyme labels within femtoliter vol- p24 molecules, resulting in a digital signal of either “active” (posi-

umes, single molecules are readily detectable by a low cost CCD tive) or “inactive” (negative) wells (Rissin and Walt, 2006). Positive

camera (Rissin et al., 2010). This simple concept is used to power a wells are counted and averaged for each array to give ‘average

standard sandwich immunoassay for p24 antigen to NAT-level sen- enzymes per bead’ (AEB), which represents the fundamental mea-

sitivity for detection of acute HIV infection. The sensitivity of the surement unit for Simoa. At higher p24 concentrations, when all

immunoassay is demonstrated by comparison to NAT and 4th gen- wells become occupied by at least one labeled p24 molecule, dig-

eration immunoassay with serial seroconversion sample sets from ital measurements transition to conventional non-digital (analog)

HIV infected individuals. The assay is standardized to quantitate measurements of total fluorescence intensity. These analog mea-

p24 protein, and correlation between p24 concentration and viral surements are converted into AEB based on the known average

RNA copies/mL by quantitative NAT is shown with HIV infected fluorescence intensity generated by single enzymes, as determined

serum. The data suggest that low cost digital immunoassay technol- by digital measurements made at low p24 concentrations. With

ogy could have important implications for HIV screening efficacy single-molecule sensitivity, concentrations of labeling reagents can

and economics. be lowered, resulting in reduced nonspecific background. This

enables high signal-to-background ratios at extremely low ana-

lyte concentrations. Because large numbers of positive wells can

2. Materials and methods be simultaneously counted on a single array, lengthy sequen-

tial bead counting is not needed for statistically powered data.

2.1. Recombinant p24 This approach makes Simoa digital immunoassays compatible

with the fast processing cadence of a high throughput automated

The recombinant p24 was obtained from Novartis Vaccines & immunoassay analyzer. The immunoassay described in this report

Diagnostics (Emoryville, CA). The HIV-1 p24 antigen is a polypep- utilizes standard low-cost reagents to demonstrate that a short

tide encoded by the p24 gag region (Steimer et al., 1986) of the diffusion-constrained signal amplification step enables thousands-

HIV-1 SF2 genome (Sanchez-Pescador et al., 1985) (231 amino fold greater sensitivity for p24 than conventional immunoassays.

acids; HIV-1 amino acid residues 139–369) and is produced in Together with nature’s amplification of 2000 p24 molecules per

L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160 155

HIV, this simple immunoassay can detect less than 100 viruses/mL,

comparable to the sensitivity of NAT for detection of acute HIV

infection.

2.5. Digital immunoassay reagents

Three reagents were developed: paramagnetic p24 cap-

ture microbeads, biotinylated detector, and a streptavidin:␤-

galactosidase (S␤G) conjugate. The capture beads were comprised

of a monoclonal anti-p24 antibody (Capricorn HIV-018-08204),

covalently attached by standard coupling chemistry to 2.7 ␮m

carboxy paramagnetic microbeads (Varian). The antibody-coated

6

beads were diluted to a concentration of 5 × 10 beads/mL in PBS

buffer with a surfactant and BSA. Biotinylated detector reagent

was comprised of a second monoclonal anti p24 antibody (Zep-

tometrix 801136), which was biotinylated using standard methods

and diluted to a concentration of 0.2 g/mL in a PBS diluent con-

taining a surfactant and goat serum, GS (PBS/GS). S G was prepared

by covalent conjugation of purified streptavidin (Thermo Scien-

tific) and ␤G (Sigma) using standard coupling chemistry. For assay,

Fig. 1. Dose–response of digital immunoassay over three logs of spiked p24 in

aliquots of a concentrated S␤G stock were diluted to 15 pM in

human plasma. Y-Axis refers to the average number of label enzymes per individ-

PBS/NCS with 1 mM MgCl2.

ual microbead (AEB) captured in each well of the array. Each data point depicts the

mean of three replicates. Standard deviations for each set of replicates are within the

circles. (a) Log-log representation of the data with 0 pg/mL calibrator omitted. (b)

2.6. Calibration Non-log depiction of the 0–0.1 pg/mL range (0 pg/mL calibrator included) showing

the low background obtained with digital quantification. Analyses of 11 calibration

curves gave a mean signal:background ratio at 0.01 pg/mL of 2.00. Statistics depict

The assay was calibrated using recombinant p24 protein diluted 2

least squares fit of all calibrators (R > 0.999). LoD was estimated as 2.8 fg/mL as

in pooled human serum (Seracare HS210). Calibrator levels were 0,

described in the text.

0.001, 0.010, 0.022, 0.100, 1.000 and 10.00 pg/mL.

2.9. Conversion between p24 units and RNA copies/mL

2.7. Immunoassay procedure

Estimates on the number of p24 units composing the HIV capsid

have ranged from 1000 to 3000 (Briggs et al., 2004; Summers et al.,

Bead-sample incubations and labeling of immunocomplexes

1992). We used an assumption of 2000 p24 units/virion, per Piatak

were conducted as described (Rissin et al., 2010), except that sam-

et al. (1993) and accepted by National Institute of Allergy and Infec-

ples were assayed without pre-dilution. In brief, the prototype

tious Disease (NIAID, 2011). HIV contains 2 copies of RNA/virion.

digital p24 assay requires 100 ␮L of sample and was performed

in three steps: analyte capture with 500,000 anti-p24 coated

microbeads for 120 min, biotinylated detector antibody binding 3. Results

for 60 min, and labeling with S G for 30 min. Assays were run in

conical 96-well ELISA plates (Axygen) using a Tecan EVO 150 for 3.1. Dose–response and limit of detection

fluidics handling and bead washing between assay steps. Serum

samples were tested with no sample pretreatment. Following assay A digital ELISA for p24 utilizing commercially available antibod-

and bead collection with a magnet, beads were loaded onto the ies and recombinant p24 for calibration was developed as described

arrays for imaging in a loading buffer comprised of PBS and 0.01% previously for other proteins (Rissin et al., 2010; Wilson et al., 2011;

Tween-20, MgCl2, and sucrose. Zetterberg et al., 2011). A representative dose–response over three

logs of spiked recombinant p24 in human serum is depicted in

Fig. 1. The assay demonstrated a highly linear response, with least

2

2.8. Array imaging squares R > 0.999. In a study of 11 calibration curves on differ-

ent days, the mean signal:background ratio (as AEB) at 0.01 pg/mL

Beads from the ELISA were loaded onto the arrays as described was 2.00 (SD 0.35). The analytical limit of detection (LoD) was esti-

(Rissin et al., 2010). Wells containing beads with labeled p24 were mated by reading the p24 concentration from the calibration curve

visualized by the hydrolysis of enzyme substrate (RGP, Invitro- at the AEB signal corresponding to 3SD above the mean signal of

gen) by ␤G into fluorescent product. Enzyme-containing wells were zero calibrator (n = 3). LoD was calculated for each of 11 calibration

imaged by fluorescence microscope fitted with a CCD camera. The runs from triplicate measurements of the zero calibrator and the

images were analyzed to determine AEB as described (Rissin et al., lowest p24-containing calibrator (0.01 pg/mL). The mean LoD was

2011). At <70% active beads relative to total beads (low p24), the 4.87 fg/mL (SD 2.89 fg/mL) (Fig. 2). Day-to-day reproducibility of a

signal output is a count of active beads corrected for a low sta- Simoa digital ELISA has recently been described in detail (Wilson

tistical probability of multiple enzymes/bead (Rissin and Walt, et al., 2011).

2006). At >70% active beads (higher p24), the probability of multiple

enzymes/bead increases, and average fluorescence of the wells is

3.2. Cultured standardized virus vs. recombinant p24

converted to AEB based on the average intensities of wells contain-

ing single enzymes determined at lower concentrations. The AEB

Cultured HIV-1 Group M, Subtype B virus was standardized for

unit thus works continuously across the digital and analog realms

RNA copies/mL and diluted in human serum as described in Section

(Rissin et al., 2011).

2. Fig. 3 depicts a log-log representation of assay dose–response to

156 L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160

0.20 b

0.10 0.08 0.1 0.06 0.04

0.02

10 10010001x10 1x10

AEB 0.02

0.01 8 a

0.01 6

0.01 4 0.01 2

0.01

0.01 0.00 8

0.00 6

0.00 4

020 040060080010001200

1000 1x104

RNA copies/mL

Fig. 3. Log-log depiction of assay response to cultured HIV-virus in human serum,

standardized for RNA copies/mL. (a) depicts low-end data down to virus-free serum

Fig. 2. Analytical LoD estimate for the digital immunoassay. A mean LoD of in non-log space. Estimated LoD was 90 copies/mL (3.6 fg/mL p24 equivalents). (b)

4.87 fg/mL (SD 2.89) was estimated from each of 11 calibration runs performed depicts data from a separate study comparing cultured virus (open circles) and p24

on different days. For comparison, LoD ranges for p24-reactive conventional antigen (closed circles).

immunoassays are depicted (11,000–70,000 fg/mL Biomerieux VIDAS p24 II, VIDAS

DUO Ultra, Abbott AxSYM HIV Combo, ARCHITECT HIV Combo) along with the LoD

of the PROCLEIX system (60 RNA copies/mL). comparison between digital immunoassay results with a quan-

titative bDNA NAT method, an immunoassay for p24, and two

4th generation HIV combo immunoassays with four sets of late-

cultured virus, and Fig. 3a highlights the low-end data down to

emerging seroconversion samples from HIV infected individuals.

virus-free serum in non-log space. Calculated analytical sensitivity

The data shown for the comparator methods were obtained from

was equivalent to 90 RNA copies/mL (or 45 viruses/mL), consistent

the sample vendor. Red represents the point in time at which the

with estimates using spiked p24 antigen. Antigen from both sources

assays first detected the HIV infection. The digital immunoassay

exhibited good agreement using an equivalency conversion based

detected the presence of p24 protein on the same blood draws that

on 2000 p24 molecules per virus (Fig. 3b), in agreement with the

the NAT method detected the presence of amplified HIV RNA. On

stoichiometry accepted by the National Institute of Allergies and

the other hand, the conventional immunoassays reactive for p24

Infectious Disease (NIAID, 2011).

did not detect the presence of antigen until one to four blood draws

later (average of 7–9 days). With seroconversion panel 9031, the

3.3. Detection of acute HIV infection Abbott tests exhibited reactivity at draw #17 (3 draws, 15 days

later, not depicted) and the Coulter p24 assay exhibited first reac-

The sensitivity of the digital immunoassay for detection of tivity at draw #18 (4 draws, 22 days later). Noteworthy is that

acute HIV infection was examined with 10 sets of seroconversion seroconversion panel 9031 comprised more frequent blood draws,

serum samples obtained from commercial sources. Table 1 depicts a and the initial reactive draw was early in the NAT-detectable phase

Table 1

Earliest detection of HIV infection.

HIV infected donor Seroconversion Days since Digital p24 Zeptometrix data

sample designation first draw immunoassay

fg/mL Chiron bDNA Coulter p24 Abbott Architect Abbott Prism

copies RNA/mL immunoassay HIV combo HIV combo

S/CO S/CO S/CO

9016-07 23

1 9016-08 27 1127.4 7473 0.32 0.21 0.20

9016-09 30 14089.9 69,101 3.36 1.41 1.11

9018-06 19

9018-07 22 45.8 304 0.09 0.14 0.07

2

9018-08 26 5067.3 15,280 0.21 0.78 0.44

9018-09 29 52692.4 193,100 1.55 5.46 4.18

9024-10 46

3 9024-11 50 1743.7 12,840 0.60 0.38 0.25

9024-12 54 Not tested >500,000 12.41 30.84 25.94

9031-13 126

9031-14 130 13.7 197 0.51 0.06 0.07

4

9031-16 137 1516.5 10,507 0.49 0.28 0.23

9031-18 152 Not tested 173,075 5.671 6.10 4.41

Mean days 57.3 57.3 66.3 64.5 64.5

L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160 157

105

Y = M0 + M1*X M0 -1.501 1 104 M1 1.1817 R 0.99464

1000

100

10 Digital immunoassay (fg/mL p24)

1

4 5 6

10 100 1000 1x10 1x10 1x10

bDNA (RNA copies/mL)

Fig. 4. Correlation between digital immunoassay and quantitative bDNA NAT

Fig. 5. Histogram of digital immunoassay results from 45 HIV-negative samples.

method. Samples tested were from four panels of seroconversion samples from HIV

Samples were tested in replicates of 3 with two reagent and calibrator batches on

infected individuals. Digital immunoassay data depicted are the mean of three repli-

different days. The dashed lines a, b, and c depict the mean AEB of the zero calibrator

cates. bDNA data are as reported by the commercial sample vendor (Zeptometrix).

(0.00559), 4SD from the HIV-negative sample population mean (0.00710), and 3SD

from the mean zero calibrator signal (0.00781) respectively. The LoD estimated at

0.00781 AEB corresponds to 5.75 fg/mL p24.

of the infection (197 RNA copies/mL). The digital p24 assay also

detected the infection in this blood draw, measuring 13.7 fg/mL

p24. First detection of this infection by conventional immunoassay

in Fig. 4 were as reported by the sample vendor, and details around

came 2–3 weeks later.

the testing were not available. The results between the two meth-

Table 2 summarizes earliest detection data across six additional

ods demonstrated linear proportionality across a 5-log range, and

sets of seroconversion samples for the digital immunoassay, three

the relationship between p24 and RNA copies/mL in this compar-

NAT methods, and two conventional immunoassay methods reac-

ison may reflect differences in standardization between the two

tive for p24. The digital immunoassay and NAT methods detected

NAT methods.

the presence of infection approximately 9–11 days earlier than the

conventional immunoassay methods. The time-to-detection for the

digital immunoassay (20.5 days) was within the range reported 3.5. HIV-negative samples

for the NAT methods (18.8–21.3 days). Taken together, these data

indicate that the digital immunoassay is able to detect acute HIV A preliminary evaluation of HIV-negative samples was con-

infection with the sensitivity of NAT. ducted on EDTA plasma from 45 presumed normal human donors.

All samples were NAT-negative for HIV RNA (PROCLEIX ULTRIO,

3.4. Correlation between p24 and quantitative NAT Novartis Diagnostics). Approximately half the samples (24) were

tested with the p24 digital immunoassay using one batch of

A comparison between the digital p24 immunoassay and a reagents and calibrators, and the remaining samples were tested

quantitative NAT method across four sets of seroconversion pan- on a separate date using a second batch of reagents and calibra-

els (Table 1) is depicted in Fig. 4. The two methods exhibited tors. The distribution of immunoassay results from all 45 samples

2

excellent correlation (R 0.99). The relationship between measured is depicted in Fig. 5. All but one sample gave an AEB signal below

p24 and RNA copies/mL reported for the bDNA method differed the mean signal for the zero calibrator (Fig. 5a). A comparison of the

by approximately 3.5-fold from that obtained by testing a stan- statistics of the HIV-negative samples with the AEB corresponding

dardized preparation of cultured virus (Fig. 3). In that study, the to the mean LoD estimated from the zero calibrators (as 3SD above

cultured virus was standardized for HIV-1 RNA/mL by the Roche the zero background) is depicted in Fig. 5b–c. The negative pop-

®

AMPLICOR HIV-1 MonitorTM Assay, v. 1.5, and the standardiza- ulation mean was over four standard deviations lower than the

tion was confirmed in the digital immunoassay by agreement with AEB corresponding to the mean LoD (5.75 fg/mL). Using either cri-

the accepted stoichiometry of p24 molecules to virus. The NAT data terion (4SD from negative sample mean or estimated LoD) as a

Table 2

Days of initial detection of HIV infection since first draw.

HIV infected Seroconversion Digital p24 Seracare data

donor panel designation immunoassay

Roche CAP/ Siemens bDNA Abbott HIV Perkin Elmer p24 Abbott Architect

CTM v1.0 NAAT NAAT RNA NAAT immunoassay HIV combo

immunoassay

5 PRB968 15 15 15 15 26 26

6 PRB956 40 40 40 40 47 47

7 PBR958 2 0 0 0 9 7

8 PBR967 0 3 7 7 17 17

9 PBR969 55 55 55 53 63 63

10 PBR972 11 0 11 3 18 18

Mean days 20.5 18.8 21.3 19.7 30.0 29.7

158 L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160

preliminary cut-off for HIV reactivity, the specificity of this limited pathogens (e.g. HIV/HBV/HCV triplex and West Nile virus), and

study was 100%. public expectation of maximum blood safety has led to the estab-

lishment of NAT for blood bank screening in higher income

countries despite its cost. This is not the case, however, for many

4. Discussion

European and developing countries (Laperche, 2005; Ekiaby et al.,

2010), where feasibility studies weigh cost effectiveness, infra-

The data presented in this report demonstrate that by isolat-

structure requirements, affordability, and disease prevalence vs.

ing single labeled immunocomplexes and confining fluorescent

less sensitive alternatives. The practicality of extending NAT pool-

product from a brief amplification step to a small volume, a

ing to clinical screening has also been explored. Stekler et al. and

simple immunoassay with a commercially available antibody

Hutchinson et al. concluded that with regular 3rd generation anti-

pair exhibited a 2000–3000-fold reduction in limit of detec-

HIV antibody testing, NAT for acute HIV infection detection makes

tion vs. conventional immunoassays reactive for p24 antigen.

sense only in higher risk settings such as sexually transmitted dis-

This sensitivity improvement, together with the natural 2000-

ease clinics (Stekler et al., 2009; Hutchinson et al., 2010).

fold amplification provided by the p24:virus stoichiometry, enable

Economics aside, timely detection and reporting represent

immunoassay sensitivity to virus comparable to NAT for detection

another challenge limiting the potential epidemiological impact of

of acute HIV infection. Preliminary analytical LoD was shown to

NAT detection of acute HIV infection. The acute phase represents

be 4.87 fg/mL, approximating 122 RNA copies/mL (∼60 virus/mL).

a period of high viremia and hyperinfectiousness, contributing

Within the sample volume of 100 ␮L utilized for the assay, this

disproportionately to the spread of the disease (Wawer et al.,

equates to detection of only six viruses. For comparison, commer-

2005; Gray et al., 2001). Turnaround time for executing a pool-

cially available 4th generation HIV combo assays exhibit LoDs of

ing strategy, testing, and deconvolution of a positive pool can delay

approximately 11–70 pg/mL (11,000–70,000 fg/mL) (Fiebig et al.,

reporting an acute infection for days, making timely intercession

2003). Across 10 sets of seroconversion samples from HIV infected

of a patient’s potential to transmit virus difficult. The time spent

individuals, the digital immunoassay matched NAT for early detec-

determining a positive result and locating the infected individ-

tion of HIV.

ual may largely negate the sensitivity advantage gained by NAT. If

A low-cost, easily deployable alternative to NAT could have

the method by which early stage infection is caught were capable

important implications for HIV screening economics and efficacy.

of a short, patient-specific turnaround of results, the reduction of

NAT economics in different screening settings is a key issue,

the detection window could have greater practical significance, as

prompting many cost-effectiveness examinations (Stramer et al.,

there would be greater likelihood of conveying acute HIV infection

2004; Marshall et al., 2004; Jackson et al., 2003; Custer et al.,

status to the patient and preventing further transmission dur-

2005; Stekler et al., 2009; Hutchinson et al., 2010; Laperche, 2005;

ing the hyperinfectious stage. Today’s automated high-sensitivity

Ekiaby et al., 2010). In blood bank screening, despite donor pool-

immunoassays typically have an assay time of an hour or less,

ing to reduce cost, NAT is cost ineffective for serology-negative

and results can be turned around while the patient waits. By

detection of acute HIV infection. In the US, estimates of cost

infusing sample with antibody-coated microbeads, even extremely

per Quality Adjusted Life Year (QALY) range from $1.5–4.3 mil-

low abundance antigen can be quickly and efficiently captured

lion unadjusted for inflation (Marshall et al., 2004; Jackson et al.,

and labeled with good ELISA-grade antibodies. This kinetic prin-

2003) which greatly exceeds normal medical standards (gen-

ciple was recently shown to apply to a very high sensitivity

erally $100,000–200,000 USD (Hutchinson et al., 2010)). Overall

Simoa digital immunoassay (Chang et al., 2012). Additionally, with

cost effectiveness ratios improve with the inclusion of additional

Fig. 6. Revised HIV screening algorithm proposed by the Center for Disease Control at the APHL HIV Diagnostics Conference, 2010.

Adapted from Rollins (2010).

L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160 159

immunoassay, virus detection does not require sample pretreat- specimens, adding $7.16/specimen to the initial immunoassay cost

ment during early stage infection when competing antibodies are (Gous et al., 2010). These studies highlight both the epidemiological

weak or absent, further enabling a short turnaround time. benefits of NAT-sensitive screening for acute HIV infection, as well

While studies on the practicality of NAT in the clinical set- as the challenges and economic pressures around NAT adoption.

ting have focused on immunoassay-negative samples, recent HIV A low cost immunoassay with the sensitivity of NAT could repre-

screening recommendations utilize NAT in a confirmatory capacity sent an opportunity to further impact the spread of transmitted and

(Owen et al., 2008; Rollins, 2010). Expanded screening is aimed transfusion-borne HIV in lower resource, high-risk areas.

at the 20% of HIV infected individuals unaware of the HIV sta- The prototype digital p24 immunoassay described in this report

tus (Marks et al., 2006), and a recent algorithm proposed in the utilizes standard low cost bead-based ELISA reagents and com-

U.S. by the Centers for Disease Control (modeled on Western Euro- mercially available antibodies. Array technology readily lends to

pean practice for the past decade) starts with a 4th generation HIV multiplexing, and from a reagent standpoint, the cost/test for a

combo immunoassay (Fig. 6). A premise of the algorithm is to uti- ‘combo’ digital p24/HIV-1/2 antibody assay would be expected to

lize 4th generation immunoassay screening due to its reactivity be comparable to conventional HIV combo immunoassays – cur-

to sufficiently elevated p24 prior to seroconversion (Hecht et al., rently one-third to one-tenth the cost of NAT (Karris et al., 2012).

2002) as well as anti-p24 immunoglobulins following seroconver- For low cost automation and deployment of digital immunoas-

sion. Fig. 6 represents inclusion of acute HIV infection detection says, Kan et al. recently reported proof of principle of low cost

capability to assay point-of-entry testing for clinical HIV screening. plastic disposable disks composed of 24 flow cells that integrate

Because HIV combo immunoassays do not discriminate between microfluidics with high-density Simoa microarrays of 216,000-fL

possible sources of a reactive result, additional discrimination sized wells (Kan et al., 2011). This array disk consumable is aimed

reflex testing is recommended. Ideally, a screening test should offer at enabling cost-effective integration of Simoa technology with

three things: high sensitivity for both acute and chronic infection, conventional automated immunoassay instrumentation perform-

low cost, and short turnaround time to facilitate patient inter- ing paramagnetic bead-based assays. More extensive validation of

vention. NAT provides the highest sensitivity for acute, but can the digital p24 immunoassay described and further development

miss chronic infection when viral load is undetectable. Laboratory of a high-throughput automated analyzer instrument is ongoing.

immunoassays and point-of-care devices offer cost and turnaround

time advantages, but can miss acute infection. 4th generation Disclosure

combo immunoassays have presented the best trade-off among

these imperatives. Although 4th generation combo immunoassays

The co-authors are employees of Quanterix Corporation.

shorten the detection window relative to serological tests, a sen-

sitivity gap remains: patients given a clean bill by 4th generation

Acknowledgments

immunoassay may actually be highly infectious. Failure to diag-

nose acute HIV infection represents a key public health problem

The authors thank Todd Campbell, Ray Meyer, Tomasz Piech,

(Pao et al., 2005; Brenner et al., 2007). Assured by apparent HIV

and Alfred Schroff for their experimental help and manuscript sug-

negative status, acutely infected individuals represent a high trans-

gestions.

mission risk and a lost opportunity to control the spread of the virus.

<10% of the 300–400 annual HIV infection diagnoses in Seattle, for

References

example, were of acute HIV infection (Stekler et al., 2009). A simple

immunoassay with rapid turnaround time that is capable NAT sen-

Assal, A., Barlet, V., Deschaseaux, M., Dupont, I., Gallian, P., Guitton, C., Morel, P.,

sitivity for acute HIV infection could represent another step toward

David, B., De Micco, P., 2009. Comparison of the analytical and operational per-

leak-proof HIV screening in a more cost and clinically effective way. formance of two viral nucleic acid test blood screening systems: Procleix Tigris

and cobas s 201. Transfusion 49, 289–300.

Because of their relative simplicity, immunoassays are more

AVERT. AIDS and HIV Information from AVERT. AVERT, Horsham, United Kingdom.

practically deployable than NAT in diverse environments. HIV

http://www.avert.org/safricastats.htm (accessed November 2011).

screening in developing countries and low-resource settings with Brenner, B.G., Roger, M., Routy, J.P., Moisi, D., Ntemgwa, M., Matte, C., Baril, J.G.,

high HIV incidence faces the issues described above, but on a larger, Thomas, R., Rouleau, D., Bruneau, J., Leblanc, R., Legault, M., Tremblay, C., Charest,

H., Wainberg, M.A., 2007. High rates of forward transmission events after

more urgent scale. NAT is a sophisticated technology with adop-

acute/early HIV-1 infection. The Journal of Infectious Diseases 195, 951–959.

tion barriers on both supplier and user sides. Although screening

Briggs, J.A., Simon, M.N., Gross, I., Kräusslich, H.G., Fuller, S.D., Vogt, V.M., John-

donated blood with NAT has been in practice since the late 90s, son, M.C., 2004. The stoichiometry of Gag protein in HIV-1. Nature Structural

& Molecular Biology 11, 672–675.

40% of worldwide blood donations remain unscreened or screened

Busch, M.P., Kleinman, S.H., Jackson, B., Stramer, S.L., Hewlett, I., Preston, S., 2000.

by less sensitive immunoassay (Global blood safety and availability

Nucleic acid amplification testing of blood donors for transfusion-transmitted

WHO fact sheet, 2009; Editorial, 2007), and 5–10% of new HIV infec- infectious diseases. Report of the Interorganizational Task Force on Nucleic Acid

Amplification Testing of Blood Donors. Transfusion 40, 143–159.

tions are transfusion-borne (Busch et al., 2005). In South Africa, the

Busch, M.P., 2007. Evolving approaches to estimate risks of transfusion-transmitted

prevalence of HIV infection in the general population is almost 11%

viral infections: incidence-window period model after ten years. In: Dax, E.M.,

(AVERT, 2011), but diagnosis of acute HIV infection is rare (Stevens Farrugia, A., Vyas, G.N. (Eds.), Advances in Transfusion Safety – vol. IV, Develop-

ments in Biologicals (Basel), vol. 127. S. Karger Publishers, Inc., Farmington, CT,

et al., 2005). Since NAT is the only methodology currently capa-

pp. 87–112.

ble of definitively diagnosing acute HIV infection, feasibility of NAT

Busch, M.P., Glynn, S.A., Stramer, S.L., Strong, D.M., Caglioti, S., Wright, D.J., Pap-

implementation in these very high incidence settings has been palardo, B., Kleinman, S.H., 2005. A new safety strategy for estimating risks of

transfusion-transmitted viral infections based on rates of detection of recently

examined (Stevens et al., 2005; Karim et al., 2007). Karim et al.

infected donors. Transfusion 45, 254–264.

demonstrated the clinical benefit of a pooling strategy for identify-

Chang, L., Rissin, D.M., Fournier, D.R., Piech, T., Patel, P.P., Wilson, D.H., Duffy, D.C.,

ing acute HIV infection in a high-risk cohort in South Africa (Karim 2012. Single molecule enzyme-linked immunosorbent assays: theoretical con-

et al., 2007). A similar study on patients attending primary health- siderations. Journal of Immunological Methods 378, 102–115.

Constantine, N.T., van der Groen, G., Belsey, E.M., Tamashiro, H., 1994. Sensitivity of

care clinics in Johannesburg showed nearly 1% of individuals were

HIV-antibody assays determined by seroconversion panels. AIDS 8, 1715–1720.

acutely infected, translating to an annual incidence rate of 12.9%

Custer, B., Busch, M.P., Marfin, A., Petersen, L., 2005. The cost-effectiveness of

(Gous et al., 2010). Gous and colleagues reported 4 cases of acute screening the U.S. blood supply for West Nile virus. Annals of Internal Medicine

143, 486–492.

HIV infection missed by 4th generation immunoassay testing of

Editorial, 2007. Improving blood safety worldwide. Lancet 370, 1879–1974.

3005 samples in a Johannesburg general hospital. The pooling strat-

Ekiaby, M.E., Lelie, N., Allain, J.P., 2010. Nucleic acid testing (NAT) in high

egy spread a $21,568 USD cost for 211 NAT results across the 3005 prevalence–low resource settings. Biologicals 38, 59–64.

160 L. Chang et al. / Journal of Virological Methods 188 (2013) 153–160

Fiebig, E.W., Wright, D.J., Rawal, B.D., Garrett, P.E., Schumacher, R.T., Peddada, L., Procleix ULTRIO Assay Product Insert. 501807-REG Rev. 3. http://www.fda.gov/

Heldebrant, C., Smith, R., Conrad, A., Kleinman, S.H., Busch, M.P., 2003. Dynamics downloads/biologicsbloodvaccines/bloodbloodproducts/approvedproducts/

of HIV viremia and antibody seroconversion in plasma donors: implications for licensedproductsblas/blooddonorscreening/infectiousdisease/ucm092120.pdf

diagnosis and staging of primary HIV infection. AIDS 17, 1871–1879. (accessed November 2011).

Global blood safety and availability facts and figures from the 2007 blood safety Rissin, D.M., Kan, C.W., Campbell, T.G., Howes, S.C., Fournier, D.R., Song, L., Piech, T.,

survey. WHO Fact Sheet #279, November 2009. Patel, P.P., Chang, L., Rivnak, A.J., Ferrell, E.P., Randall, J.D., Provuncher, G.K., Walt,

Gous, N., Scott, L., Perovic, O., Venter, F., Stevens, W., 2010. Should South Africa be D.R., Duffy, D.C., 2010. Single-molecule enzyme-linked immunosorbent assay

performing nucleic acid testing on HIV enzyme-linked immunosorbent assay- detects serum proteins at subfemtomolar concentrations. Nature Biotechnology

negative samples? Journal of Clinical Microbiology 48, 3407–3409. 28, 595–599.

Gray, R.H., Wawer, M.J., Brookmeyer, R., Sewankambo, N.K., Serwadda, D., Wabwire- Rissin, D.M., Fournier, D.R., Piech, T., Kan, C.W., Campbell, T.G., Song, L., Chang, L.,

Mangen, F., Lutalo, T., Li, X., vanCott, T., Quinn, T.C., 2001. Probability of HIV- Rivnak, A.J., Patel, P.P., Provuncher, G.K., Ferrell, E.P., Howes, S., Pink, B.A., Min-

1 transmission per coital act in monogamous, heterosexual, HIV-1-discordant nehan, K.A., Wilson, D.H., Duffy, D.C., 2011. Simultaneous detection of single

couples in Rakai, Uganda. Lancet 357, 1149–1153. molecules and singulated ensembles of molecules enables immunoassays with

Hecht, F.M., Busch, M.P., Rawal, B., Webb, M., Rosenberg, E., Swanson, M., Chesney, broad dynamic range. Analytical Chemistry 83, 2279–2285.

M., Anderson, J., Levy, J., Kahn, J.O., 2002. Use of laboratory tests and clinical Rissin, D., Walt, D., 2006. Digital concentration readout of single enzyme molecules

symptoms for identification of primary HIV infection. AIDS 16, 1119–1129. using femtoliter arrays and Poisson statistics. Nano Letters 6, 520–523.

Hutchinson, A.B., Patel, P., Sansom, S.L., Farnham, P.G., Sullivan, T.J., Bennett, B., Rollins, G., 2010. What’s the best testing strategy for HIV infection? Clinical Labora-

Kerndt, P.R., Bolan, R.K., Heffelfinger, J.D., Prabhu, V.S., Branson, B.M., 2010. tory News 36, 1–11.

Cost-effectiveness of pooled nucleic acid amplification testing for acute HIV Sanchez-Pescador, R., Power, M.D., Barr, P.J., Steimer, K.S., Stempien, M.M., Brown-

infection after third-generation HIV antibody screening and rapid testing in the Shimer, S.L., Gee, W.W., Renard, A., Randolph, A., Levy, J.A., 1985. Nucleotide

United States: a comparison of three public health settings. PLoS Medicine 7, sequence and expression of an AIDS-associated (ARV-2). Science 227,

e1000342. 484–492.

Jackson, B.R., Busch, M.P., Stramer, S.L., AuBuchon, J.P., 2003. The cost-effectiveness Steimer, K.S., Puma, J.P., Power, M.D., Powers, M.A., George-Nascimento, C., Stephans,

of NAT for HIV, HCV, and HBV in whole-blood donations. Transfusion 43, J.C., Levy, J.A., Sanchez-Pescador, R., Luciw, P.A., Barr, P.J., 1986. Differential

721–729. antibody responses of individuals infected with AIDS-associated

Kan, C.W., Rivnak, A.J., Campbell, T.G., Piech, T., Rissin, D.M., Mösl, M., Peterc¸ a, A., surveyed using the viral core antigen p25gag expressed in bacteria. Virology

Niederberger, H-P., Minnehan, K.A., Purvish, P.P., Ferrell, E.P., Meyer, R.E., Chang, 150, 283–290.

L., Wilson, D.H., Fournier, D.R., Duffy, D.C., 2011. Isolation and detection of single Stekler, J.D., Swenson, P.D., Coombs, R.W., Dragavon, J., Thomas, K.K., Brennan, C.A.,

molecules on paramagnetic beads using sequential fluid flows in micro fabri- Devare, S.G., Wood, R.W., Golden, M.R., 2009. HIV testing in a high-incidence

cated polymer array assemblies. Lab on a Chip 12, 977–985. population: is antibody testing alone good enough? Clinical Infectious Diseases

Karim, S.S., Mlisana, K., Kharsany, A.B., Williamson, C., Baxter, C., Karim, Q.A., 2007. 49, 444–453.

Utilizing nucleic acid amplification to identify acute HIV infection. AIDS 21, Stevens, W., Akkers, E., Myers, M., Motloung, T., Pilcher, C., Venter, F., 2005. High

653–655. prevalence of undetected, acute HIV infection in a South African primary health

Karris, M.Y., Anderson, C.M., Morris, S.R., Smith, D.M., Little, S.J., 2012. Cost savings care clinic. In: IAS Conf. HIV Pathol. Treatmt., Rio de Janeiro, Brazil (Abstr.

associated with testing of antibodies, antigens, and nucleic acids fordiagnosis of MoOa0108).

acute HIV infection. Journal of Clinical Microbiology 50, 1874–1878. Stramer, S.L., Glynn, S.A., Kleinman, S.H., Strong, D.M., Caglioti, S., Wright, D.J., Dodd,

Kontio, S., 1991. Sensitive one-step enzyme immunoassay for HIV-1 p24 antigen in R.Y., Busch, M.P., 2004. Detection of HIV-1 and HCV infections among antibody-

human blood specimens and cell culture supernatants. Journal of Immunological negative blood donors by nucleic acid-amplification testing. The New England

Methods 139, 257–263. Journal of Medicine 351, 760–768.

Laperche, S., 2005. Editorial. Blood safety and nucleic acid testing in Europe. Euro Summers, M.F., Henderson, L.E., Chance, M.R., Bess, J.W., South, T.L., Blake, P.R., Sagi,

Surveillance 10 (2). I., Perez-Alvarado, G., Sowder, R.C., Hare, D.R., 1992. Nucleocapsid zinc fingers

Marcel, M., Grèzeb, M., Bailly, A., Izopet, J., 2011. Analytical sensitivity of four HIV detected in retroviruses: EXAFS studies of intact viruses and the solution-state

combined antigen/antibody assays using the p24 WHO standard. Journal of Clin- structure of the nucleocapsid protein from HIV-1. Protein Science 1, 563–574.

ical Virology 50, 57–60. Ukkonen, P., Korpela, J., Suni, J., Hedman, K., 1988. Inactivation of human immu-

Marks, G., Crepaz, N., Janssen, R.S., 2006. Estimating sexual transmission of HIV from nodeficiency virus in serum specimens as a safety measure for diagnostic

persons aware and unaware that they are infected with the virus in the USA. AIDS immunoassays. European Journal of Clinical Microbiology & Infectious Diseases

20, 1447–1450. 4, 518–523.

Marshall, D.A., Kleinman, S.H., Wong, J.B., AuBuchon, J.P., Grima, D.T., Kulin, N.A., Wawer, M.J., Gray, R.H., Sewankambo, N.K., Serwadda, D., Li, X., Laeyendecker,

Weinstein, M.C., 2004. Cost-effectiveness of nucleic acid test screening of vol- O., Kiwanuka, N., Kigozi, G., Kiddugavu, M., Lutalo, T., Nalugoda, F., Wabwire-

unteer blood donations for hepatitis B, and human immunodeficiency Mangen, F., Meehan, M.P., Quinn, T.C., 2005. Rates of HIV-1 transmission per

virus in the United States. Vox Sanguinis 86, 28–40. coital act, by stage of HIV-1 infection, in Rakai, Uganda. The Journal of Infectious

Mullis, K.B., Faloona, F.A., Scharf, S., Saiki, R.K., Horn, G., Erlich, H.A., 1986. Specific Diseases 191, 1403–1409.

enzymatic amplification of DNA in vitro: the polymerase chain reaction. Cold Weber, B., Fall, E.H., Berger, A., Doerr, H.W., 1998. Reduction of diagnostic window

Spring Harbor Symposia on Quantitative Biology 51, 263–273. by new fourth-generation human immunodeficiency virus screening assays.

National Institute of Allergies and Infectious Disease. Structure of HIV. Journal of Clinical Microbiology 36, 2235–2239.

http://www.niaid.nih.gov/topics/HIVAIDS/Understanding/Biology/pages/ Westreich, D.J., Hudgens, M.G., Fiscus, S.A., Pilcher, C.D., 2008. Optimizing screening

structure.aspx (accessed November 2011). for acute human immunodeficiency virus infection with pooled nucleic acid

Owen, S.M., Yang, C., Spira, T., Ou, C.Y., Pau, C.P., Parekh, B.S., Candal, D., Kuehl, D., amplification tests. Journal of Clinical Microbiology 46, 1785–1792.

Kennedy, M.S., Rudolph, D., Luo, W., Delatorre, N., Masciotra, S., Kalish, M.L., Wilson, D.H., Hanlon, D.W., Provuncher, G.K., Chang, L., Song, L., Patel, P.P., Fer-

Cowart, F., Barnett, T., Lal, R., McDougal, J.S., 2008. Alternative algorithms for rell, E.P., Lepor, H., Partin, A.W., Chan, D.W., Sokoll, L.J., Cheli, C.D., Thiel,

human immunodeficiency virus infection diagnosis using tests that are licensed R.P., Fournier, D.R., Duffy, D.C., 2011. Fifth-generation digital immunoassay for

in the United States. Journal of Clinical Microbiology 46, 1588–1595. prostate-specific antigen by single molecule array technology. Clinical Chem-

Pao, D., Fisher, M., Hué, S., Dean, G., Murphy, G., Cane, P.A., Sabin, C.A., Pillay, D., 2005. istry 57, 1712–1721.

Transmission of HIV-1 during primary infection: relationship to sexual risk and Zetterberg, H., Mörtberg, E., Song, L., Chang, L., Provuncher, G.K., Patel, P.P., Fer-

sexually transmitted infections. AIDS 19, 85–90. rell, E., Fournier, D.R., Kan, C.W., Campbell, T.G., Meyer, R., Rivnak, A.J., Pink,

Piatak, M., Saag, M.S., Yang, L.C., Clark, S.J., Kappes, J.C., Luk, K.C., Hahn, B.H., Shaw, B.A., Minnehan, K.A., Piech, T., Rissin, D.M., Duffy, D.C., Rubertsson, S., Wilson,

G.M., Lifson, J.D., 1993. High levels of HIV-1 in plasma during all stages of infec- D.H., Blennow, K., 2011. Hypoxia due to cardiac arrest induces a time-dependent

tion determined by competitive PCR. Science 259, 1749–1754. increase in serum amyloid ␤ levels in humans. PLoS ONE 6, e28263.