EPA-650/2-74-044


April  1974
Environmental Protection Technology  Series

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                                   EPA-650/2-74-044
ISOLATION  OF  HAYFEYER  ANTIGENS
   FROM  SHORT RAGWEED  POLLEN
                       by

                   Robert M. Flora

            VVorthington Biochemical Corporation
               Freehold, New Jersey 07728
                Contract No. 68-02-0566
               Program Element No. 1AA010
            EPA Project Officer.  Eva Wittgenstein

             Chemistry and Physics Laboratory
           National Environmental Research Center
         Research Triangle Park, North Carolina  27711
                    Prepared for

          OFFICE OF RESEARCH AND DEVELOPMENT
         U.S. ENVIRONMENTAL PROTECTION AGENCY
               WASHINGTON, D.C.  20460

                     April 1974

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This report has been reviewed by the Environmental Protection Agency
and approved for publication.  Approval does not signify that the
contents necessarily reflect the views and policies of the Agency,
nor does mention of trade names or commercial products constitute
endorsement or recommendation for use.
                                  11

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                               -1-
Abstract:  Short ragweed pollen has been processed to yield antigens
associated with hayfever allergy. Antigens K and Ra3 have not been
processed to purity but were collected and retained as by-products
during the preparation of antigen E.  Antigen E has been examined
by physical and immunological means and found to  meet high
standards of purity.

Introduction:   A fixed-price contract for $40, 000 was established
for Worthington Biochemical Corporation to use accepted,  published
methods for isolating antigens  from short ragweed pollen.  The
basic process uses extraction, salt  precipitation, and column
chromatography.   Process  steps are monitored by testing samples
with antisera specific for the individual antigens and by  polyacrylamide
disc gel electrophoresis.  Worthington Biochemical Corporation has
operated this process several times prior to  this run for the Environ-
mental  Protection Agency,  and was  fortunate.in encountering no
difficulties this time.  The  process  behaved normally and gave yields
which fell into  the anticipated range.

Materials and Methods

Short ragweed  pollen,  Lot 18-15-72, consisting of 20 x  1 kg bottles
was provided by EPA to the contractor in November of 1972.  Informa-
tion about the handling or drying of  the pollen or of its crop year  was
not provided.   The pollen was held in cold storage from the time  of
arrival until processed.

All chemicals utilized were of reagent grade and suppliers were
the following;  tris (hydroxymethyl)  amino methane (THAM),  Fisher
Scientific; ethyl ether,  sodium chloride,  sodium hydroxide and
hydrochloric acid, J. T. Baker; ammonium sulfate, J.  T. Baker
or Mallinckrodt.   Column chromatographic media were: Sephadex
G-25 and G-100,  and DEAE Sephadex A-50,  Pharmacia  Fine Chemicals;
Celite  545,  Johns-Manville.

Antisera were  from  the following sources; anti-antigen  E,  lot 92008A,
and anti-whole ragweed, 1022P,  N.  I. H. Research Reagents distributed
by the  National Institute of  Allergy and Infectious Diseases; anti-antigen
K,  2KA, Dr. T. P.  King; anti-antigen Ra3,  2KA,  Dr. L. Goodfriend.

Immunodiffusion plates were from Kallestad  Laboratories.  Immuno-
electrophoresis was performed in a thin film agarose medium using
films and equipment supplied by  Analytical Chemists Incorporated.
Disc electrophoresis in polyacrylamide gel and micro-Kjeldahl analyses
were, conducted in a standard manner by the Worthington Quality  Control
Department.

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                                 -2-
Isolation Procedures
   *
Purification of antigen E was based upon methods published by T.  P.
King and co-workers.  ^« ^« •*'

The substitution of DEAE Sephadex A- 50 for TEAE cellulose and later
DEAE cellulose was a significant difference between the process
described by King and the procedure followed by Worthington.  One
deviation from the technical proposal was required; it was
not feasible to  carry out recycling gel filtration over Sephadex G-100
because available optical monitoring units could not handle high
absorbancies encountered at the scale of this work.  An additional,
final step not in the original technical proposal was incorporated as
an ammonium sulfate gradient/Celite 545 chromatographic procedure
more recently  described by
All chromatographic work was monitored by appropriate immunological
techniques, absorbancy at 280nm,  and in latter stages by disc gel
electrophoresis.

1.   Defatting of pollen with ethyl ether

     The 20 kg of pollen was  suspended in 60 liters of ethyl ether,
     slurried manually then mechanically over a period of 30 to 60
     minutes followed  by transfer to filtration apparatus for solvent
     removal.  The pollen cake was resuspended in 50 liters of
     ethyl ether and stirred mechanically for 30 minutes,  again
     transferred to filter funnels and the cakes washed extensively
     with ether until the filtrate was pale yellow.  The washed pollen
     was spread and air-dried until the odor of ether was no longer
     detectable.  The defatting operation was conducted at ambient
     temperature; all subsequent operations were at 4°C.

2.   Extraction of the  pollen  antigens

     The ether extracted dried pollen was suspended in 100 liters of
     cold deionized water and stirred mechanically overnight (16-18 hrs).
     The aqueous extract was collected by filtration, the  pH adjusted to
     7. 0 with 1. ON NH40H and brought to cold saturation with ammonium
     sulfate, 690g /liter.   The pollen cake was resuspended in 80 liters
     of cold deionized  water, stirred overnight, filtered, and the second
     aqueous extract treated  as described above.  Twice  more the pollen
     cake was extracted with water, 60 liters each time,  the extracts
     neutralized and brought  to saturation with ammonium sulfate.

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                                -3-
2. x Extraction of the pollen antigens (cont'd. )

    The aqueous extractible, ammonium sulfate precipitable material
    was collected by centrifugation, dissolved in water, all four
    extracts combined,  pH adjusted to 7. 0, and the solution clarified.

3.  Column I;  Sephadex G-25

    The 9500 ml solution of crude ragweed antigens was desalted and
    freed of considerable pigment by passage over a 40 x 100 cm bed
    of Sephadex G-25 equilibrated and eluted with 0. 025M Tris-Cl
    buffer, pH  7. 9.  All fractions containing appreciable protein and
    preceding the elution of the ammonium sulfate were pooled.

4.  Columns Ha and lib; DEAE Sephadex A-50

    The crude ragweed  protein after desalting was applied directly
    to two in-parallel columns,  21. 5 x 100 cm. , of DEAE Sephadex
    A-50 equilibrated with 0. 025M Tris-Cl buffer, pH 7.9, followed
    by washing with the equilibrating buffer until a volume of approx-
    imately 60 liters for each column had  been collected.   Antigens
    Ra3, Ra5, and BPAP were eluted with the equilibration buffer
    and these pools were precipitated with ammonium sulfate, dissolved,
    dialyzed, and lyophilized as the side fractions from columns Ila and
    lib. Antigen E and  accompanying antigen K were eluted by applica-
    tion of a non-linear gradient derived from 24 liters,  each vessel, of
    equilibration buffer and 0. 05M Tris-Cl, 0.25m NaCl, pH 7. 9.
    Fractions containing antigens  E and K were combined, precipitated
    with ammonium sulfate, collected and redissolved  in minimal
    volume and dialyzed against column III buffer.

5.  Columns Ilia.  Illb and IIIc; Sephadex G-100

    The antigen E and K solution in 0. 05m Tris-Cl, 0. 4M ammonium
    sulfate,  pH 7.4, (approximately 1200 ml) was  applied to a column
    bed, 21. 5 x  100 cm. of Sephadex G-100 packed with the same
    buffer.  Fractions containing antigen E essentially free of antigen K
    were pooled and held.  Fractions containing antigen E  contaminated
    with antigen K were pooled, concentrated, and passed over an
    identical column, Illb, of Sephadex G-100.  Again antigen E was
    pooled and held separately from the antigen E and K containing
    fractions.   A third  pass over G-100,  column IIIc, was performed.
    Main pools  of antigen E from columns Ilia, Illb and IIIc were

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                                -4-
5. x Columns Ilia, Illb and IIIc; Sephadex G-100 (cont'd. )

    combined.  Leading fractions  to the main E pools were combined,
    precipitated with  ammonium sulfate,  redissolved,  dialyzed,
    clarified and lyophilized as  "leading side fractions", columns
    Illabc.   Trailing side fractions,  not subject to repeat passage
    over the Sephadex G-100 columns,  were treated  in a similar
    manner and were lyophilized as "trailing side fractions"
    columns Illabc and should contain the bulk  of antigen K.

6.  Column IV;  DEAE Sephadex A-50

    The combined antigen E pool from the previous step  was
    dialyzed against equilibration buffer,  0. 025M Tris-Cl,  pH 7. 9,
    prior to application to a 10 x 100 cm column bed of DEAE
    Sephadex A-50.  Elution was effected by ^developing a linear
    gradient,  10 liters each vessel, of the equilibrating buffer and
    0.025M Tris-Cl, 0.4m Nad, pH 7. 9.  Fractions containing
    the substitative portion of the  antigen E were combined.  Leading
    and trailing side pools were also made; these were dialyzed and
    lyophilized as side fractions,  column IV.

7.  Column V; Cel'le 545/Ammonium Sulfate Gradient

    On the finding that antigen E from column  IV had not attained the
    desired state of purity,  an additional  chromatographic procedure
    was employed.  A trial  run with 5 per cent of the material
    demonstrated that the ammonium sulfate gradient procedure would
    further purify antigen E.  The bulk of antigen E from column IV
    in 0. 05M Tris-Cl, pH 8. 0,  was precipitated by 0. 7 saturation of
    ammonium sulfate in the presence of Celite 545.  The resulting
    protein precipitate-Celite slurry was packed into a 5 x 50 cm
    column over a small plug of Celite. The column was washed with
    0. 7 saturation of  ammonium sulfate following by a gradient,
    2 liters each vessel of decreasing ammonium sulfate between the
    limits of the original conditions down to the simple buffer, 0. 05M
    Tris-Cl,  pH 8.0.

    The pool of fractions containing the antigen E from column V was
    extensively dialyzed until free of salt, including  the small amount
    from the trial run and the entire  E pool concentrated approximately
    five-fold in an Amicon ultrafilter device with a 10, 000 molecular
    weight cut-off membrane to a final antigen E concentration of 12 mg/
    ml.

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                                -5-
  x Antigen K obtained from column V was precipitated by ammonium
    sulfate,  redissolved,  dialyzed and lyophilized, and provided as
    one of the by-product materials.

    Antigen E obtained in bulk from the processing described was
    subjected to final contractor testing as described below.  On
    the determination that it met the required specifications of
    quality it was prepared for final lyophilization,  dispensed
    into brown 50cc bottles and lyophilized.  Fifty-four bottles
    containing 100 mg. of antigen E were delivered and one
    bottle, of approximately 50 mg, was retained by the contractor.

Results;

Characterization of Antigen E,  Lot  53H382
                                           «
I   Immunological (Micro-Ouchterlony) Analysis

    A.   Employing specific anti-antigen E serum:
         Antigen E at concentrations ranging from 1200 to
         12 ug/ml vs. anti-antigen E serum (#92008)
         exhibits a single  continuous precipitin line
         establishing the  identity of the material as
         antigen E.  Refer to  (Plate I).

    B.   Employing an anti-whole ragweed serum:
         Antigen E at concentrations ranging from 1200  to
         12 ug/ml vs. anti-whole ragweed serum (#1022P)
         exhibits again a  single continuous precipitin line
         (Plate II).  At the highest level, 1200 ug/ml,
         another diffuse line is visible.  Based upon the
         level of the sample which shows the contaminant
         line relative to the concentration of the  sample
         applied it is estimated that the contaminant does
         not exceed a level of 40 ug/ml or 3. 3% of the
         antigen E.  The  indication  from immunoelectro-
         phoresis is that  this  contaminant is antigen K.
         It should be  noted that the specification  calls
         for the antigen concentration to be over the
         range 10 to 200 ug/ml.   Where concentrations of
         the antigen from 12 to 400  ug/ml are used, a
         contaminant line is not observed.

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                               -6-
II   Immunoelectrophoresis

    Antigen E exhibits a triphasic precipitin line against both
    anti-antigen E and anti-whole ragweed sera, correlating
    with the three bands visible in disc polyacrylamide gel
    electrophoresis. At the level of 1Z mg/ml a distinct arc
    is observed against the anti-whole ragweed serum and a
    somewhat less distinct line against anti-antigen K serum.
    At 4 mg/ml the  contaminant line is considerably diminished
    against the anti-whole ragweed serum and almost imper-
    ceptible vs. the anti-antigen K serum. Negative  results
    were obtained with anti-antigen Ra3 serum.

Ill  Ultraviolet Spectrum

    A scan (Fig. 2)  of the  antigen E over the wavelength range
    400 to  2ZO nm indicates the preparation possesses an
    adsorptivity which is typical of protein in general, z°"'250 =
    2.40,  and characteristic of antigen E  in particular.  Freedom
    from significant pigment contamination is  indicated by the
    adsorbancy,  280'320 = 22.7.  Both ratios  exceed the contract
    specifications, which are, 280/250^-  2.2 and 280/320^ 15.

IV  Disc Electrophoresis in Polyacrylamide Gel

    Antigen E obtained exhibits three bands on disc gel electro-
    phoresis.  A densitometric scan (Fig.  3) indicates the bands
    to be equi-spaced and  in the approximate ratio of 15:60:25.
    Since antigen E  may contain as many as four electrophoretically
    separable components, designated A-D, this finding is compatible
    with published reports.  Preparations conducted over the past 18
    months at Worthington Biochemical Corporation have been
    observed to favor the isolation of more of the A-form and lesser
    of the  D-form of antigen E.  Photo 1.

    Comparison of lot 53H382 with lot 1KA delivered previously to
    the EPA reflects a greater degree of antigen E homogeneity
    as evidenced by freedom from the uppermost band (not in the
    equi-spaced pattern) and a lesser amount of leading edge
    smearing.   Photo 2.

    Admixture of a  standard antigen E preparation,  XPTA, and
    lot 53H382 can be reconciled with the  following.   The standard,
    primarily forms B and C,  with a trace of  D band,  co-electro-
    phoresised with lot 53H382,  some A but mainly B,  and C bands,
    gives the pattern of four visible bands as expected, A, mainly
    B,  C,  and a trace of D.   Photo 3.

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                               -7-
IV  Disc Electrophoresis in Polyacrylamide Gel (cont'd. )
   X
    The side fraction (Column V; llOmg) containing antigen K is shown
    compared to a standard antigen K, 3BA.   Photo 4.

V  Nitrogen Content

    Micro-Kjeldahl analysis has determined that the nitrogen content
    of the antigen E is 15.4%, within the specification range of  15-17%.

Description of Fractions Obtained as By-products,  Lot 53H38Z

Antigens K and Ra3 were not obtained in high states  of purity and
thus have  not been subject to critical analysis.  The following
fractions obtained as by-products of the Antigen E preparation
have been delivered as lyophilized bulks of pooled fractions and
are listed with a general description.

1.  Side Fractions - Column II;  354 g.  Known to contain antigen
    Ra3 and suspected to contain in addition antigens Ra5 and BPAR.
    Relatively crude.

2.  Leading Side  Fractions - Columns Ilia, IHb and IIIc;
    8.6 g.  Contains impure antigen E.

3.  Trailing Side Fractions - Columns Ilia, Illb and IIIc;
    6.5 g.  Contains impure antigen K.

4.  Leading Side  Fractions - Column IV;  3. 0 g.  Contains
    some impure antigen E.

5.  Trailing Side Fractions - Column IV;  2.4 g.  Contains
    some antigen E and antigen K.

6.  Side Fractions - Column V;  110 mg.   Contains  relatively
    pure antigen K.

References

1.  King, T. P., Norman, P. S.,  and Connell,  J.  T.  Biochem. ^,
    458 (1964).
2.  King, T. P., Norman, P. S., and Lichtenstein, L. M.  Biochem.
    6,  1992 (1967).
3.  King, T. P.  Biochem.  U_,  367 (1972).

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        .":.-)..•, ,•.-.;, ••-' •,.'.--..-.  •>"-•., ', ' '.-.- -•- . .• ..  •'

                       PLATE  I

Mlcro-Ouchterlony analysis employing anti-
antigen  E  serum,  92008A, vs. antigen E,
53H382.

Center well:   Antt-antlgen E serum

Wells  1-5=   1200, ^00,  120, AO,  and 12 pg/ml
antigen  E.
                                                     PLATE  I I

                               Mlcro-Ouchterlony analysis  employing anti-
                               whole  ragweed serum, 1022P,  vs.  antigen E,
                               53H382.

                               Center well:   Ant I -whole ragweed serum

                               Wells  1-5:   1200, 1»00, 120,  1*0,  and 12 yg/ml
                               antigen  E
.'••.-.••• •:.!-!!f:;!i!:-i!fi-!-!:;-!-!::;:'.: Hfijigjigp!!*?!.
                                                                           ^^

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                       FIGURE I
          THIN FILM AGAROSE IMMUNOELECTROPHORESIS
                    D


                    E
                                                2


                                                3


                                                1*



                                                5


                                                6
                    H
Wells A-D:  Antigen E, 53H382, 12 mg/ml ; 2 microliters
            In each wel 1 .

Wells E-H:  Antigen E, 53H382, k mg/ml; 2 microliters
            In each wel 1 .
Trough 1 and 7:

Trough 2 and 6:

Trough 3 and $:

Trough k       :
                   Anti-antigen Ra3 serum, lot 2KA

                   Anti -antigen K serum, lot 2KA

                   Anti-whole ragweed serum, #1022P

                   Anti-antigen E serum, #92008A
(Approximately ^tO microliters serum applied to trough)

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                                                           FIGURE  2
    1.0
    0,8 -
    0.6 -
CO
I
s
IE
    0.4 -
    0.2 -
                                                     ANTIGEN  E, 53x382

                                                     GARY  KOBEL 15 SCAN
                                                     0.80  KG/I! IN 0.01 H TRIS-HCL, PH 7.9
           2"0
280
        320

WAVELENGTH, NANOMETERS
360
                                                                                                                   400

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                                                   FIGURE  3
I
                                                              AIITIGEM E, 53H382
                                                            DENSIOflETRIC SCAN OF
                                                           POLYACRYLAMIDE DISC GEL
                                                              ELECTROPKORESIS

                                                     FULL SCALE:  2.5 OPTICAL DENSITY
                                                     RATE OF SCAN:  2 INCHES EQUALS
                                                     1 CM OF DISC GEL TRAVEL
                                                     PROTEIN:  40 KICROGRAMS
                                           INCHES

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             Antigen E, l/>t H382
     80 ug   40 ug    24 ug   12 ug
                           «.

                PHOTO  1

Antigen E, 53H382, polyacrylamide  disc  gel
electrophorests, pH 8.6.  Mlcrograms  antigen
E applied, left to right, 80,  40,  2^  and  12.
           PHOTO 2

Antigen E, lot  1KA and  lot 53H382
        ;/•'; •'::;?"::".''''.'                  ....,..,
 ' •                '             ' '        '   •

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                            MS 8"^-
Antigen E, left, standard, lot 01A,'right, lot
53H382, and centeV, mixture of 01A and 53H382.
Antigen K, 53H382, a by-product of Column V,
compared with an antigen K standard.
      • .': '.r. .1 '^
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                                   TECHNICAL REPORT DATA
                            (Please read Instructions on the reverse before completing}
1. REPORT NO.
                                                           3. RECIPIENT'S ACCESSION-NO.
4. TITLE AND SUBTITLE
                                                           5. REPORT DATE
  Isolation of Hayfever  Antigens from Short Ragweed
  Pollen
              April 26. 1974
             6. PERFORMING ORGANIZATION CODE
7. AUTHOR(S)

  Robert M. Flora
                                                           8. PERFORMING ORGANIZATION REPORT NO.
9. PERFORMING ORGANIZATION NAME AND ADDRESS

  Worthington  Biochemical Corporation
                                                           10. PROGRAM ELEMENT NO.
              1AA010
             11. CONTRACT/GRANT NO.
                                                            68-02-0566
12. SPONSORING AGENCY NAME AND ADDRESS
  Environmental  Protection Agency
  Chemistry  and  Physics Laboratory
  Research Triangle Park, M. C. 27711
             13. TYPE OF REPORT AND PERIOD COVERED
              Final.  Nov.  72 - Mar. 73
             14. SPONSORING AGENCY CODE
15. SUPPLEMENTARY NOTES
16. ABSTRACT
  Contract  objectives were to isolate antigens  from short ragweed  pollen for use as
  standard  reference materials in the study  of  aeroallergens and atmospheric proteins.

  Antigen E was  prepared in pure form while  both of the minor antigens,  K and Ra3,
  were not  processed to purity.  Twenty  kilogram of pollen were defatted and extracted
  with water.  The antigens were then isolated  by salt precipitation  and column
  chromatography.   The process was monitored with electrophoresis  and immune precipitin
  tests.  The  final  bulk of antigen E obtained  was tested for purity  and lyophilized
  before shipment.  Total  yield was 5.4g of  pure antigen E and approximately 374g of
  the side  fractions containing K and Ra3.   Micro-Ouchterlony tests of antigen E gave
  a single  precipitin line with anti-antigen E  serum.  Immunoelectrophoresis resulted
  in a triphasic precipitin band corresponding  to the three bands  obtained in poly-
  acrylamide gel  electrophoresis.  UV absorption scans of the antigen E  showed a
  typical protein  peak at 230 nm wavelength, and micro-Kjeldahl analysts gave a
  nitrogen  content of 15.4%.  Antigens K and Ra3 were not critically  tested.  All test
  results were compatible with published reports and indicated that the  antigen E
  prepared  meets a high degree of purity.
17.
                               KEY WORDS AND DOCUMENT ANALYSIS
                  DESCRIPTORS
                                              b.IDENTIFIERS/OPEN ENDED TERMS
                          c.  COSATI Field/Group
  Hayfever
  Ragweed  Pollen
 Aeroallergens
 Column Chromatography
 Isolation of Antigen E
18. DISTRIBUTION STATEMENT
            •
  Release  Unlimited
19. SECURITY CLASS (This Report)
 Unclassified
                                                                         21. NO. OF PAGES
                                              20. SECURITY CLASS (This page I

                                               Unclassified
                                                                         22. PRICE
EPA Form 2220-1 (9-73)

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