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Diurnal variation of tree pollen in the air in finland

Markku Käpylä

To cite this article: Markku Käpylä (1984) Diurnal variation of tree pollen in the air in finland,
Grana, 23:3, 167-176, DOI: 10.1080/00173138409427712

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Grana 23: 167-176, 1984

Diurnal variation of tree pollen in the air in Finland

hiARKKU KAPYLA

Kfipyla, hf. 1984. Diurnal variation of tree pollen .in the air in Finland. - Grana
23: 167-176, 1984. Uppsala 8 November 1984. ISSN 0017-3134.
The pollen w a s collected with Burkard spore traps inTurku and Jyvaskyla, in southern and
central Finland. Selected days of high concentration were analysed for hour to hour
concentrations. The diurnal variation of pollen concentration w a s variable in Alnrrs,
Betuln, Jrrriiperrts and Pinrts, while Picen, Poprrlrts, Qirercrts, Snlix and Ulrriirs had regular
daytime maxima och nightly minima. The pollen concentrations were closely correlated
with increasing temperatures and decreasing relative humidities. A significant relationship
with wind speed i t a s also found, except in A l m s and Berrrla. The most significant
relationships with wind speed were found in Snlix and Qirercrrs, the pollen of which tends
t o remain in lumps. Generally, the bigger the pollen grains are, the more their concentra-
tions are dependent on wind speed. Pinrrs and Picen seem to have a threshold wind speed
about 4 m / s above which concentrations become higher.
hlnrkkri Kiipyli, Depnrtriietit of Biology, UniLversity of JpiisXylii. SF-40100 Jyiiskylii 10,
Finlurid.
(hlntirtscripr receiced I I hlny 1983, revised cersiori nccepred 2 Jnrirrnry 1984)

Few published measurements are available about
the diurnal variation of tree pollen in the air. Diur-
nal patterns of pollen emission are usually studied
by placing a pollen trap in the middle of the source
vegetation. Distant from the sources the diurnal
pattern of pollen occurrence is affected by the pol-
len emission rhythm of the source as well as the
movement of the air masses, which makes the si-
tuation very complex. In this paper the diurnal
variation of tree pollen in the air in Finland is
studied in places fairly distant from local sources.
The aim of the study is to get preliminary knowl-
edge on the main factors affecting diurnal vari-
ation. The results can be applied to the research
and treatment of pollen allergies.
hlATEKIAL AND METHODS
Locutions of the study
The collecting localities were the same as in the previous
publication concerning the diurnal variation of non-arbo-
real pollen in the air (KapylS 1981). One locality was the
roof of the Science Building of the University of Turku I5
m above ground (cf. hlakinen & Rantio-Lehtimiki 1979).
Another trap \ i s situated in Jyvaskyla Airport, 1.5 m
above ground in an open place (cf. KBpyla et al. 1980).
The weather data are from standard meteorological meas-
urements done in the weather stations of Jyvaskyla and
Turku Airports.
Collecting nnd corrnring pollen
Pollen collecting w a s done with Burkard spore traps in the
usual itay. The details of the procedure have been dc-
scribed by Rantio-Lehtimaki & hlakinen (1981).Selected
days from the peak pollen periods were taken in this
study. The slides of these days were studied microscopi-
cally with 24 transverse traverses corresponding to every
full hour (cf. Kapyla & Penttinen 1981).
The selected days are the following: Alnrts: Turku:
13-18.1V.1976; 15, 18, 20, 24 and 25.1V.1978. Berrrlnr
Turku: 18-24.V.1977; 16-27.V.1978. Jrrnipcrrrs: Turku:
11-14, 16.VI.1977; 3-9.V1.1978; JyvBskyla Airport (used
only in Fig. 3): 14, 16-17.VI.1977, 5-7, 9-10, 13 and
16.V1.1978.'Picen: Turku: 27-28.V.1978; Jyviskyla Air-
port: 30.V-I.VI.1978. Pinrrs: Turku: 11-14, 16-17.VI.
1977; 3-9.V1.1978; Jyvaskyla Airport (used only in Fig.
3): 1 1 , 14, 16-17.V1.1977, 5-7, 9-10, 12-14 and 16.VI.
1978. Poprtlrrs (only in Fig. 2): Turku: 9-13.V.1976;
17-20.V.1978. Qirercirs: Turku: 9-12.V1.1977; 31.V.-
I.V1 and 3.V1.1978; 31.V-2.V1.1979. Sn1i.r: Turku:
12.V.1976; 16, 19,27and 28.V.1978;17.V. and 2.V1.1979.
Only in Fig. 3: 1 1 and 13.V.1976; 31.V.-I.V1.1978. Ul-
rtrris: Turku: 12.V.1976; 16-19.V.1978; 16-17.V.1979.
Only in Fig. 3: I 1 and 13.V.1976.
The material does not allow comparison of the two

Crana 23
168 Af. Kapyla

Frequency RESULTS AND DISCUSSION

J-

The diurnal patterns of pollen occurrence

were variable (Fig. 2). That is why the distri-
150- bution of the peaks along different hours of the day
and night and their sizes is more informative (Fig.
3). The effect of some weather factors was also
studied. The weather factors were so closely corre-
100-
lated that their effects are very difticult to separate
with multivariate methods. One way analyses were
performed on temperature (Fig. 4). relative humid-
50-
ity (Fig. 5) and wind speed (Fig. 6). The variation in
the material is very big but the general pattern
probably reflects real relationships, not necessarily
causal.
0 -r-
0
Na of pollen grains
Ainus (Alder)
In the Turku area this genus is represented almost
exclusively by the species Ainus glutinosa. Ainus
incana is very rare in that area.
The diurnal pattern of the occurrence of alder
pollen in the air is very irregular. The peaks OC-
curred at any time but the highest peaks were found

h,
during the night. During the periods of hour to hour
studies of pollen occurrence the temperature varied
from -4 to 15°C and the diurnal peaks coincided
L.0
with temperatures -3 to +7"C and the relative
humidity varied from 31 to 100%. The diurnal
Log N o of pollen grains
peaks coincided with relative humidities 46100%.
Fig. I. Frequency distribution of the pollen data of Befitla There were significant correlations and F-values
in arithmetic (above) and logarithmic scale (below). between pollen amounts and temperature and rela-
tive humidity but not with wind speed. The wind
speeds ranged from 0 to 13.5 d s and the diurnal
peaks were in wind speeds 26.5 d s , except once
(18.IV.1976) a second peak in a wind velocity of
localities. To exclude the effects of possible differences,
only the observations from Turku were used in analyzing
13.5 d s .
the influence of weather factors (except in Picea). During rain only low concentrations were found
The daily mean pollen counts of Turku can be found in and no clear decline of concentrations were found
the Finnish Pollen Bulletin and of Jyvaskyla in Kapyla et because of rain. Especially snowfall seemed to
al. (1980). have no effect.
Siarisrics The studies of Rempe (1937), Stix & Grosse-
The pollen data are about log-normally distributed (Fig. Brauckmann (1970) and Janzon et al. (1977) showed
1). That is why logarithmic transformations were used in highest frequencies on the average in the afternoon.
most cases. Single-factor analyses of variance were per-
formed on the effects of temperature, relative humidity
and wind speed on pollen concentrations. The differences
of the means of the factor levels were compared with Befirla (Birch)
Student-Newman-Keuls test (Sokal & Rohlf 1969: 244). Different species of Betirln were not separated in
When a linear relationship was found, correlation was the pollen counts because of difficulties in the iden-
also calculated. The significance levels were shown by the
following symbols: *p<O.OS, **p<O.OI, ***p<O.OOl. tification.
Other details are given in the figures. There is no regularity in the diurnal pattern of

Grana 23
 Diiirtial variatioti of tree pollen in Fitilarid 169
Alnus
*/.

0 1

Betula hpulus
a./

0 1

Juniperus puercus

'0 1

- . . . . . . . . . . . ,
Pice0 Soh

101

Pinus Ulmus

'$1
Fig. 2. The mean occurrence of pollen
grains at different hours of the day and
l
0 a
6 12 m
18 24 night as percentages of the total.
hours hours

occurrence of birch pollen. The peaks may occur tween these species should be studied in pure
about as often in any time of the day and night, stands, which are big enough.
except perhaps early in the morning. Also the aver-
aged concentrations were about the same through- Jiiniperiis cornmiinis (Juniper)
out the day and night. In most days the pollen The peak pollen occurrences may be at any time of
occurrence was closely correlated with increasing the day and night, but mostly during daytime and
temperatures and decreasing relative humidities. In near midday. This same is reflected in the averaged
some cases the concentrations became higher to- diurnal pollen occurrence curve. The variation is
wards the night, provided that the relative humidity fairly closely correlated with increasing tempera-
did not become very high and the night temperature tures and decreasing relative humidities. With wind
was relatively high (more than 10°C). The effect of speed there was no correlation, but F-value was
wind speed was not statistically significant. The statistically significant. In comparing the means of
small material of Stix & Grosse-Brauckmann (1970) the wind levels (Fig. 6), only the second level d i t
showed similar trends as in the present study. fered significantly from the two levels surrounding
Sarvas (1955) found about noon peaks for both B. it.
pendiila and B. piibescens, but the sampler he used
is not reliable in studying diurnal variation. He also Picen abies (Norway Spruce)
found a slight difference in the hours of peak pollen Only five days of high occurrence of spruce pollen
occurrence between these species. The possible was available for the study. According to this mate-
differences of the timing of pollen emission be- rial high occurrence seems to be during day and a

Cram 23
170 M. Kiipylii
P.g/rn’
nzll ALNUS
0 0

O0
0

100
;J , , ”.o, , , , . ,
n=26 PINUS
0

.o0 O 8
.O 0 0 1000
0 co 0
1000: 0

, -
100 OUERCUS

JUNIPERUS

0 0 100

n-10 SALIX

’cool 0

1 0 0 1
100
].. , ,, ; ,oo;n. , ,
Fig. 3. The sizes and occurrences of the
pollen peaks during different hours of the
day and night. The days with maximum
n=5 PlCEA
n=l2 ULHUS concentrations less than 100 pollen
0 0 grainslm’ and the days with no clear peak
0 0
0 are excluded. Double peaks are both in-
cluded, circles are observations from Turku

100 LJ 100 , . ;.
6 12 I’e
, ,

burs
,
li
and squares from JyvSskyll Airport, Fin-
land.

low concentration during night. According to Sar-
vas (1955) the diurnal peak was in the morning
between approximately 9.00 and 10.00 hours.
High pollen occurrence seems to coincide usually
with high temperature, low relative humidity and
fairly high wind speed. The means of the first three
wind speed levels (Fig. 6) differ significantly from
the means of the levels of high wind speeds. This
indicates a threshold wind speed around 4 d s . A
threshold might also b e with relative humidity.
Piriirs sylvestris (Scots pine)
The averaged concentration curve shows a mini-
mum early in the morning and a period of high
pollen occurrence from about noon to midnight.
The peaks are fairly evenly distributed from about
10 a.m. to midnight. The pollen occurrence was
closely correlated with increasing temperatures and
decreasing relative humidities. In days with moder-
ate temperatures (max. 17-24°C) the peak usually
coincided approximately with the maximum tem-
perature and minimum humidity. In very dry and
warm days (max. 25°C or mo%) the time of the
peak pollen occurrence was usually delayed until a
moderate humidity and temperature was reached
again. T h e mean time of the peak pollen occurrence
was significantly later in days with high maximuni
temperatures (Fig. 7). T h e reasons for this are ob-
scure.
The pollen concentrations were higher in higher
wind speeds, indicating a threshold around 4 m / s
(cf. Picea). These results differ greatly from the
observations of Sarvas (1955, 1962). H e found a
definite maximum at about noon and very little
pollen during the night. T h e type of pollen trap
Sarvas used does not work well in low wind speeds,
which probably explains the discrepancy. Other
data in the literature contain very limited material.
Scamoni (1938) found that the times of daily maxi-
ma varied. P e r s o n (1955) found a maximum at

Grana 23
 Diirrrtnl unriatiorl of tree pollert iri Firilnrtd I7 1

.r
10000
ALNUS PlNUS
F=62*** F = 28.9***
r=O.36*** r=063***

looo/

n = 26L 21
10 7 '
20 30 0 10 20

BETULA QUERCUS

iltf
F=17,6*** looo1 F = 16.2***

i
r = o.L~***

1000:

I
100

0
73
- 10 68 44 35
i0
I
L1
I

20

JUNIPERUS SALlX

I:
F = 12.9*** F :13.5***
r = 0.43***

l100
oo]

lL
10

1000
n.288

10 20
30

k
PlCEA ULHUS
F = 21.1*** F = 12.5***
r = 067***
-
-
- --

1
1007
--
-
--
10: 10

.
I.

33 17 1e 2620 26 n=lZO 30 20 8 n=168

1 . I
- I
0 10 30 'C
Temperature
Fig. 4. The relationship betiyeen temperature and pollen The number of obsenztions is shown at the base of each
concentration. Geometric means (the columns) and geo- column. F is the variance ratio and r is the coefficient of
metric standard deviations (the segments of line) are used. correlation.
 172 hi. h'apyla

r!
PINUS
F = 29.3***

!I
r .-065***

10

lo^ zf !
n=26C 15 3750 L3 50 63 6 8
1 10
10000 ° ~ 100
1 0%

10 01
BETULA QUERCUS
F = 18 5 * * * F = 23.7***
r I -0.51 ** * r = -0.57***

100

10 0

looo1
92 705055 5L 56 38

JUNIPERUS
F =5.9***
loo 50

SALIX
ia
F=8.0***
100%

r ;-030*** +**
r = -0. ~9

t
T

10

0
15
-
!I
1i.
15 55 LC 100
10

! o l 1 0

10001
n=168

T
ULMUS
F=16.2***
0 ./.

10

n -168
50 100 ./.[ol 1 0
50
Relative humidity
100 */.
Fig. 5 . The relationship between relative humidity and
pollen concentration. Explanations as in Fig. 4.

Grana 23
 Diiirnal variatioti of tree pollett it1 Fittlnttd 173

PINUS
F=5.2***

13 n = 312
T 10
i 0 4 i rnls
PUERCUS
F = 0.7
F = 1L.5***
r = -0.06
T

100
I
l 10
o o 0 c 99
1
- 8 0 4

r 7
JUNIPERUS SALIX
F = IS** F =11.1***
r i0.06 r=0.51***

,,&t
1
100

10

1 0
1 n.168
S rnls

T ULMUS
F=88***
r=O.61*** T 1

10
t
, 38 ,I 1; 7
3
-n = 168
L mls
Wind speed
Fig. 6. The relationship between wind speed and pollen
concentration. Explanations as in Fig. 4.
Grana 23

I
* I
I
meon = l5.L I meon.21.1 3.7
t i
s d e v = 3.5 I sdev.3.5 p<O.W5
20 I) *C Max temp
. <so0
500 -999
a 1093-1999
0 >?a00
poiien groins / m3
Fig. 7. The time and level of the pine pollen peak in days
with different maximum temperatures. Days with precipi-
tation more than 5 mm are omitted. The groups above and
below 25°C were compared with Student's /-test. The
regression line, regression equation and the correlation
coeffecient ( r ) are also shown.
noon and Stix and Grosse-Brauckmann (1970) early
in the afternoon.
Popitlits (Aspen and poplar)
Except the fairly common native forest tree Popir-
Iris Ireniiila. this group also contains imported pop-
lars found in parks of Turku.
T h e diurnal rhythm seems t o be very clear with a
maximum at about noon and a midnight minimum.
The material was too small for studying the effects
of weather conditions or the variation in pollen
peak hours.
Qirercirs robirr (Oak)
Oak is rare as a native and fairly common as a
planted park tree in Turku area. In the averaged
diurnal rhythm there a r e peaks at about noon and
late afternoon, and a minimum late in the night.
There is a similar double distribution of pollen
peaks. 'The pollen concentrations were the bigger.
the higher the wind speeds were. T h e pollen con-
centrations were bigger in higher temperatures and
lower relative humidities. Both figures indicate that
there might b e a threshold value for either o r both
of the weather factors.
Salix (Willows)
The averaged curve shows a broad daytime maxi-
mum and a minimum during night. The peaks are
distributed fairly evenly from 10 to 19 o'clock. Gen-
Gram 23
erally the concentrations are bigger in higher tern-
peratures and lower relativc humidities, but con-
centrations became lower in temperatures above
21°C. The concentrations increase very much with
increasing wind velocity. This is understandable
because the pollen grains of insect-pollinated wil-
lows tend to stick together and need strong agita-
tion to become airborne. T h e seemingly lowering
effect of high temperatures o n pollen concentra-
tions is probably due t o the fact that wind velocities
are lower in high temperatures (Fig. 8).
Ultiiirs (Elm)
hlost of thc elm trees found in Turku area belong to
the species Ultiiiis glabro.
The pollen peaks occurred during daytime from G
t o 19 o'clock, but the highest peaks were late in the
afternoon. With all the weather factors studied,
there seems to b e an optimum, on either side of
which concentrations diminish. T h e small material
makes the conclusions uncertain.
DISCUSSION
The diurnal rhythm of pollen occurrence of trees is
much more irregular than that of herbs and grasses
(cf. Kapyla 1981). Once started their anthesis thus
seems to proceed more independently o n prevailing
weather conditions, which makes their flowering
more liable to failure. But trees are long-lived and
that is why it is not necessary to have a successful
seed yield every year.
In many trees (Altirrs, Berirla, Jiiniperirs, Pitius)
high pollen concentrations occur during both day
and night and it is usual that the concentrations stay
for about the same fairly long periods. In other
trees (Picea, Popiiliis, Qiierciis, Salix, Uhiirs)
Wtnd speed
m/s
G1
Temperot ure
Fig. 8 . The wind speed in different temperatures (data
concerning Salk). The means are indicated by columns
and the standard deviations by the segments of line.

"1
F - v a l u e w i t h w i n d speed
,
,
_- \
,,' O o u e r c u s
I
I ,
S i z e o f t h e pollen groln
Fig. 9. The dependence of pollen concentrations on wind
speed compared to the size of the pollen grains (embedded
in gl ycerine-gclatinc).
there is a regular daytime maximum and nightly
minimum.
The diurnal variation of pollen concentration
does not necessarily reflect only the pollen liber-
ation rhythm. One possibility is that the pollen in
days with strong thermal convection and turbu-
lence goes high up in the air and falls down during
evening and night. This phenomenon might explain
part of the high evening and night concentrations
(cf. Rempe 1937). This phenomenon should be
studied. If this is true, night pollen should be on the
average smaller in size (cf. Rempe 1937) and its
viability lower.
The effect of weather factors on hour to hour
pollen concentrations is difficult to study because
the weather factors are closely correlated. Further-
more, the successive observations are not inde-
pendent but autocorrelated. This excludes the
orthodox use of most of the statistical tests. These
can, as in this study, be calculated, but the conclu-
sions must be cautious.
The aerial pollen concentrations are dependent
on pollen shedding, take-up of pollen into the air
and its transport by wind and thermal convection.
Kain means disturbance to this series of events.
These phases are differently affected by weather
conditions. The opening of anthers is probably
caused by drying. So it is understandable that in
high relative humidities pollen concentrations were
clearly lower. About as significant correlations or
F-values were obtained with temperature than with
relative humidity. That could be explained by their
Dirrrtiol vorintiori of tree pollcti irz Fitilririd 175
close correlation, but swelling of catkins and the
growth of filaments are probably also affected by
temperature. The rupture of the anthers might also
be affected by enzymatic breakdown (Stanley &
Linskens 1974:24), which means that this phase
might also be temperature-dependent. The last
phases of anthesis are different in different trees.
The effect of weather conditions on each phase
should be studied experimentally in climate cham-
bers.
Pollen release and its staying airborne is promot-
ed by air movements. When the F-value with wind
speed of each species is plotted against the size of
their pollen grains (Fig. 9), the species form three
groups. Within each group the F-value is dependent
on the size of pollen grains. The most significant
relationships between wind speed and pollen con-
centrations were found in willows (Snlix) and oak
(Qrrercrrs). Willows are insect-pollinated and their
sticky pollen tends to remain in lumps, and so need
strong agitation to become and stay airborne. Qrrcr-
C I I S , which has insect-pollinated relatives, has re-
tained some features of entomophily, e.g. the fairly
rough surface of the pollen grain. The lumping of
pollen in Qrrercrrs is high among wind-pollinated
species (Andersen 1970). The second group is
formed by normal wind-pollinated trees. Ultrzirs dif-
fers somewhat from the group with its comparative-
ly high F-value. This is in accordance with its com-
paratively high clumping of pollen (Andersen 1970).
The third group is formed by Pitiris and Piceo, the
pollen grains of which have air sacs.
The performance of the Burkard spore trap is
probably affected by wind speed and particle size,
which weakens the conclusions of this study. With
a slightly different version, the Hirst spore
trap, some wind tunnel experiments have been
made. According to Hirst (1952) the efficiency has
always been below 100%, but Ogden et al. (1974)
found that the efficiency in high air speeds was
more than 100%. Both found a minimum perfor-
mance of about 60-70% in moderate air speeds.
However, the performance in outdoor air is differ-
ent, because of the big size of the trap and its
sluggish response to rapidly changing wind direc-
tions. hlay et al. (1976) found a clearly lower sam-
pling efficiency for large liquid droplets in outdoor
experiments than the previously mentioned authors
for pollen and spores in wind tunnels. In the same
outdoor experiments no trend wind speed was
found. Thus we might think that the expected in-
Grana 23
176 hf. Kapyla

crease of the efficiency in high wind speeds is com-
pensated by the increased tilting effect due to high-
er turbulence. The conclusion is that it is best not to
use any correction coefficients with wind speed.
Furthermore, the F-values do not differ much if the
correction coefficients from the Hirst’s (1952)
curve are applied.
ACKNOWLEDGEMENTS
I would like to express my gratitude to hlrs Ritva Kupias
in Turku and hlr Heikki Veijola in Jyvaskyla, who did the
practical pollen collection and made the preparations. I
would also like to thank hlrs Ritva Kupias and hlrs Pirkko
Lehtinen who did the microscopical counting and hlr
hlatti Tikkanen and hlrs Liisa Roning-Pynndnen who
made the drawings. This study was financed by the Finn-
ish Academy of Sciences.
des nliitenstaubes durch die Luftstromungen. -.
Planta 27:93-147.
Sarvas, R. 1955. Investigations into the flowering and
seed quality of forest trees. - Comm. Inst. Forest.
Fenn. 45,47: 1-37.
Sarvas, R. 1%2. Investigations on the flowering and seed
crop of Pinus silvestris. - Comm. Inst. Forest. Fenn.
53,541 1-198.
Scamoni, A. 1938. Uber Eintritt und Verlauf der mannli-
chen Kiefernbliite. - Z. Forst- und Jagdwesen
70: 28S315.
Sokal, R. R. & Rohlf, F. J. 1%9. Biometry. - San
Francisco.
Stanley, R. G. & Linskens, €1. F. 1974. Pollen: Biology,
biochemistry, management. - Springer-Verlag. Ber-
lin, Heidelberg.
Stix, E. & Grosse-Brauckmann, G . 1970. Der Pollen- und
Sporengehalt der Luft und seine tages- und jahreszeit-
lichen SchMankungen unter mittel-europaischen Ver-
hlltnissen. - Flora 159: 1-37.

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Ogden, E. C., Raynor. G. S., Hayes, J. V., Lewis, D. hi.
& Haines, J. H. 1974. hlanual for sampling airborne
pollen. - IIafner Press, New York.
Persson, A. 1955. Frequenzen von Kiefernpollen in
Siidschweden 1953 und 1954. - Z. Forslgenetik
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