2. Bradfield, E. G. 1977. Extraction of calcium fractions from plant
material. Comm. Soil Sci. and Plant Anal. 8:563-572.
3. Epstein, E. 1972. Mineral nutrition of plants: Principles and per
spectives. Wiley, New York.
4. Hartmann, H. T., A. Tombesi, and J. Whisler. 1970. Promotion
of ethylene evolution and fruit abscission in the olive by 2-chloroethanephosphonic acid and cycloheximide. J. Amer. Soc. Hort.
Sci. 95:635-640.
5. Hoagland, D. R. and D. I. Arnon. 1950. The water culture method
for growing plants without soil. Univ. o f Calif. Cir. 347.
6. Klein, I., E. Epstein, S. Lavee, and Y. Ben-Tal. 1978. Environmental
factors affecting ethephon release in olive. Scientia Hort. 9:21-30.
7. Poovaiah, B. W. and A. C. Leopold. 1973. Inhibition of abscission
•by calcium. Plant Physiol. 51:848-851.
8. Shaw, R. 1969. The influence of 2-chloroethane phosphonic acid
(Ethrel) on maturation, ethylene production, and perianth dehis
cence. Proc. Assoc. Southern Agr. Workers Conf 66:203.
9. Sterry, J. R. 1973. The uses and potential of CEPA in fruit crops.
Acta Hort. 34:489-495.
J. Amer. Soc. Hort. Sci. 105(1):37—42. 1980.
A Computer Simulation to Maximize Asparagus Yield1
Em m ett P. Lampert, Donn T. Johnson, Art W. Tai, Glen Kilpatrick, Randy A. Antosiak,
Philip H. Crowley, and Erik D. G oodm an2
D epartm ents o f E ntom ology and Electrical Engineering & System s Science, Michigan State
University, East Lansing, M I 48824
A d d itio n a l in d ex words. A sparagus officinalis, harvest strategy, photosynthesis, plant growth
A b stra ct. A computer simulation of asparagus growth is developed and used to evaluate the effects of various
harvest strategies on short and long term commercial yield of asparagus (A sparagus o fficin a lis L.). At present
asparagus is harvested until the canners stop buying, usually in the 3rd or 4th week of June in southern Michigan,
purchase generally being terminated by the reduction of spear diameter (whips), increase in fiber content of the
spears or opening of the bracts. The simulation shows that this stragety is economically optimal for any single
year; however, if the grower terminates the harvest every year on June 1, then the average yearly yields are
significantly greater than those derived from the previous strategy. Skipping strategies, in which the grower skips
a harvest every nth year (2nd, 3rd, or 4th), produced significantly lower 15 year average yields than either of the
other 2 strageties, but produced significantly greater yields per plant.
Methods
Asparagus growth is dependent on light, temperature, nu
trients, water and other biotic and abiotic factors (12, 13, 16,
4, 2, 15, 20, 10, 3, and 18). In addition, age and sex are im
portant factors in determining the yield of a single plant (21,
7, 15, and 11), but lack of consistent information prevents
their incorporation into the model at this time. Because of
these constraints, the model was constructed and used to
evaluate the qualitative rather than the quantitative effects
of harvest strategies on plant yield. Consequently, the model
simulates the growth of a typical 7 year old plant of indeter
minate sex having 7 bud sites.
The structure of the model can be broken down into 3
periods: preharvest, harvest and postharvest (Fig. 1).
Preharvest. During this portion of the simulation, the user
has the option to change default parameter values, to select
a harvest strategy and to determine the number of plants to
simulate and the length of simulation in years (Fig. 2). The
subroutine WEATHER is called here and when executed reads
in the current years maximum and minimum daily temperature,
calculates average daily temperature and constructs arrays of
minimum daily temperature and average daily temperature for
use by the rest of the subroutines.
Harvest. This portion of the model is composed of the
subroutine HARVEST, which carries out all harvest activity
from spear emergence to termination of harvest (Fig. 2).
Although the first emergence of spears is of primary impor
tance when calculating asparagus yield, due to the loss in yield
iReceived for publication March 19, 1979. Journal Article No. 7939
Michigan Agr. Expt. Sta. This research was partially supported by EPA
grant No. R802547 “Bionomics and Management of Soil Arthropod
Pests” and No. R803785020 “Utilization of Pest Ecosystem Models in
Pest Management Programs”.
The cost of publishing this paper was defrayed in part by the payment of
page charges. Under postal regulations, this paper must therefore be
hereby marked advertisement solely to indicate this fact.
2\Ve extend our thanks to Dr. Dean L. Haynes, Department of Ento
mology, Michigan State University, for making available the resources
to complete this manuscript. We gratefully acknowledge the data and
direction provided by Drs. Hugh Price and Alan Putnam, Department of
Horticulture at Michigan State University . We would also like to acknowl
edge other members of the Entomology department. To Kenneth Dimoff
for the graphics software package used in Fig. 3 to 6, and Claudia Klepsteen for Fig. 1 and 2, we thank you.
^Rajzer, C. J. 1975. A study of asparagus buds: dominance and influence
on spear size. A special class report for Horticulture 330, Michigan
State University.
4silver Mills. Shelby, Michigan. 1975 Asparagus Grade Guide.
Fig. 1. Functional block diagram of asparagus simulation.
Fig. 2. Flow chart of asparagus simulation.
from freezing, little work has addressed this problem. LeCompte
and Blumenfield (8) found increases in predictability of first
spear emergence using degree days (1) over calendar date but
reported “considerable error” even using degree days. Since
this was the case, first spear emergence in our simulation begins
on a user defined date (May 1 was used as default first emer
gence). After the emergence of the first spear, the distribution
of primary spear growth activity from the other bud sites
greatly influences yield. Data by Rajzer3 indicate no dominance
or significant order of spear initiation between the bud sites.
Further, his data indicated a uniform distribution of primary
spear growth activity over about a 12 day period. Therefore,
in our simulation a random number generator was used to
generate a uniform distribution of primary spear growth ac
tivity over a 12 day period (subroutine EMERGE). Since
replicate runs differ in these initial dates of primary spear
growth activity, they represent to some degree the observed
variability among individual plants.
After initialization of primary bud growth at the 7 bud
sites, the subroutine YIELD is accessed. This subroutine is
responsible for growth and harvest of spears. When executed,
YIELD calls the subroutine DIAM and spear diam is calculated.
Energy for growth of spears must come directly from the
carbohydrates stored in the root. Tiedjens (22) states that
spear production and size are dependent on these stored carbo
hydrates. Also, Norton (14) found highly significant positive
correlations between total cross sectional area of ferns in the
fall and total cross sectional area of spears the following spring,
while Ellison and Scheer (4) found similar correlations as well
as a significant correlation between stalk index per plant in the
fall and spear diam the following spring. These results indicate
a relationship between spear diam and carbohydrate reserves.
Since no data were available on spear diam through a season and
carbohydrates, we were forced to look at % marketable spears.
This was a decreasing sigmoid curve (H. Price, personal com
munication). We assumed that a Michaelis-Menten relationship
existed between spear diam (D) and carbohydrate reserves
above a minimum level (Cm), and found the following equation
to be sufficiently descriptive:
D = Dm(C - CmVCDk + C - Cm),
[1]
where
Dm = maximum spear diameter,
C = current carbohydrate reserves,
Cm = minimum level of carbohydrate reserves for spear
production, and
Dk = Michaelis-Menten parameter controlling form of
equation.
Table 1. Default or initial parameter values for equations in text.
Parameter
Equation
number
Initial/default
value
C
1
282.2 g
Cm
1
168.5 g
Dk
1
Dm
1
28.0 cm
2.8
.906
7.62 cm
d
Lr
3
3
dc
Lu
4
4
fe = fl
5
.20
Kw
5
81.0 cm
R
6
.45 g/day
I
7
.40 cal
cm'2 min"1
Pi
7
.898
a
8
82.903
b2
ft
8
8
8
.043 g/cm'3
22.86 cm
8.140
.064
.0073
Source
Scott, et al. (16)
Table 14, p. 30.
Scott, et al. (16)
Table 14, p. 30.
Curve fitting to data
from Dr. Hugh Price.
Field observations.
Laboratory measurements.
Segerlind (17), and
field observations.
Scott, et al. (16) p. 29.
7.62 cm suprasurface
remains and 5.24 cm sub
soil remains. Takatori,
et al. (20).
Curve fitting to field
observation.
Curve fitting to field
observation.
Curve fitting to field
observation.
Extrapolation of data by
Downton and
TorOkfalvy (3).
Integrated proportion of
day where Intensity >
.40 cal cm'2 min'1.
Multiple regression to
data by Sawada, et al.,
(15), Table 1.
Same as above.
Same as above.
Curve fitting to field
observation.
Default or initial values for these parameters and their source
can be found in Table 1.
If the spears are not whips (diameter < .5 cm) or the carbo
hydrate reserves are above the minimal amount required for
spear production, then harvest continues and spears are grown
(Fig. 2).
Within the subroutine MOERDYKE, Julian date (IDATE)
is incremented and spears are grown. Spear growth, a function
of temperature and spear height, is calculated by the following
equation as presented by Moerdyke (10):
G = 3.9092 + .3162 H + .6379 T,
[2]
where
G = spear growth in cm/day,
H = spear height in cm, and
T = temperature in °C.
If Julian date is less than the user defined last date of harvest
or branching has not occurred, then the spears are grown and
yield calculated by the subroutine YIELD. Working (23) has
shown that branching is a function of temperature. He found
when average daily temperature was 24° or 40°C, branching
occurred from 20 to 25 cm and 6 to 8 cm, respectively. Inter
polating his data to a minimum harvest height of 11 cm, an
average daily temperature of 26° caused branching in our
simulation and terminated harvest.
The subroutine YIELD calculates both the asparagus yield
to the grower and the loss of carbohydrates to the storage root.
For a spear to be harvested, it must be greater than 18.6 cm in
length. Any spear shorter than 18.6 cm can safely be allowed
to grow for another day. In Michigan, asparagus is harvested
either by hand or with a sled. Segerlind (17) has shown a
sharp reduction in force required to cut a spear as you go from
2.5 to 10 cm from the spear base, also the cutting blades in
sleds are generally 6 to 8 cm above ground. Therefore, in our
model a 7.6 cm portion of the spear remains after harvest.
For a given spear of harvestable length, its wt (Yg) can be
calculated by:
Yg =(D/2)2 7 r(L -L r) ( 0 ( d ) ,
[3]
where
D
L
Lr
f
=
=
=
=
spear diameter (cm),
spear length above ground (cm),
remaining spear portion after harvest (cm),
correction factor = spear volume/cylinder volume,
and
d = density of spear (g/cm3).
Default or initial values for these parameters and their
source can be found in Table 1. After each harvest, the carbo
hydrate content in the harvested spear is computed and sub
tracted from the carbohydrate reserves. Either harvest or freez
ing terminate spear growth and contribute to carbohydrate
depletion. The loss of carbohydrates (Cj) from the root reserves
used in production of the spear is calculated by:
c, = (D/2 )2 7r [(L -
Lr) (f)
+L J
(d) (d,),
[4]
where
Cj = loss of carbohydrates,
Lu = unharvested length of spear, and
dc = density of carbohydrates in g/g spear fresh wt.
Default or initial values for these parameters and their
source can be found in Table 1. It is assumed there is no resorp
tion of carbohydrates by the storage root.
Postharvest. Since the yield of asparagus is a function of
carbohydrate reserves (14, 22, 4, and 15) and these reserves
are produced by the previous years ferns, it is important in
a simulation to include postharvest activity. In our simulation,
the subroutine CARB models this activity.
When executed, CARB increments the Julian date, calcu
lates daily photoperiod, grows the asparagus fern and assimilates
carbohydrates. The amount of carbohydrates produced and
stored in the root depends on spear diameter at the end of the
season (14 and 4), light intensity (cal cm-2 min'1), photoperiod,
temperature and weight of photosynthetically active tissue (15).
In the simulation, the basal diameter of the spear is used
as the diameter for the fern, which in turn is used to calculate
max leaf weight. By weighing all leaves on 6 year old plants,
Sawada et al. (15) found an average fresh weight of leaves of
630 g and 497 g for staminate and pistillate plants, respectively.
These were averaged for our simulation and the following
equation used to predict max leaf wt per fern:
Wm = fj D/(K* + D),
[5]
where
fj = parameter controlling maximum value of equation,
D = spear diameter (cm), and
Kw = Michaelis-Menten parameter controlling the form
of the equation.
Due to the lack of data, these parameters and the exact
equation form could not be determ ined. Therefore, a MichaelisMenten relationship was assumed and the 2 parameters adjusted
to produce the desired range of output. Default or initial values
for these parameters are listed in Table 1.
Leaf weight growth is assumed to behave logistically (9) as a
function of leaf weight and time and described by:
dw
= RW (Wm
dt
W)/Wm,
[ 6]
I— CNJ
where
0
MODEL
*
OBSERVED
R = intrinsic rate of growth, and
W = current weight of fern leaves.
The default value for R is listed in Table 1. Scott et al. (16)
observed a reduction in carbohydrates of 100 g per plant as a
result of fern production over a 28 day period. In our simula
tion, no attempt was made to represent this decline. Instead,
each plants’ carbohydrate reserve is depleted by 100 g over
a 28 day period to account for the loss due to fern production.
Data by Downton and Torokfalvy (3) indicates that as
irradiance increases the rate of increase in the photo synthetic
activity decreases. This implies that photosynthesis becomes
light saturated. Since photosynthesis was still increasing in
the range of their experiment, we selected .40 cal cm'2 min'1
as the irradiance at which photosynthesis saturates (2.75 times
reported range). Therefore, total cal (K) available for photo
synthesis were calculated by:
K = PIPi,
[7]
where
P = daily photoperiod in minutes,
I = light saturation irradiance (cal cm'2 min'1), and
Pi = proportion of area under normal solar intensity
curve where value > I.
Default values of I and p* are shown in Table 1. The amount
of carbohydrates assimilated (C) is in turn calculated from the
following equation:
C(t + 1) = [fT T(a + biP + b2 K) (W)] + C(t)
[8]
where
fT = curve parameter,
T = Degree days > 5.6°C,
a+fyP+bzK = multiple regression for CH20 assimulation
per 5 g leaf weight, and
W = weight of fern leaves.
See Table 1 for values and sources of parameters.
Carbohydrates continued to be assimilated by the ferns
until death or until the storage roots capacity for carbohydrates
is reached. There are 2 causes of fern death in our simulation:
1) an average daily temperature below —1.1°C will freeze the
fern, or 2) the grower chops the fern. In Michigan, growers
frequently chop the asparagus ferns in the fall to prevent
excess accumulation of snow which retards spring field access.
Since asparagus root growth from year to year is not well
documented, it was necessary to set an upper limit to the
amount of carbohydrates stored in the roots. Morse (12) re
ported that up to 56% of the asparagus roots’ dry weight was
carbohydrates and Scott et al. (16) found from 36% to 43%
in a variety of treatments. Shelton (18) examining the storage
of soluble storage carbohydrates in 3 year old plants in Michi
gan, found a range of % from 20% to 74% under a variety of
lengths of harvest. Due to this variety in %, we allowed a max
storage level of 350 g/plant; which allowed for a 25% increase
in storage (16). In multiple year runs, overwinter carbohydrate
losses due to respiration must be examined. The model a c co u n ts
for overwintering respiration loss of carbohydrates by multi
plying the final fall carbohydrate reserves by 0.932, this allows
for 6.8% of the reserves to be used for overwinter survival.
Although no actual data on asparagus was found, the value of
0.932 came from unpublished data on winter loss in another
perennial, common goldenrod (P. A. Werner, personal com
munication).
Certain other factors that may be important to asparagus
Fig. 3. Comparison of total carbohydrates in the asparagus crown. Ob
served results (Shelton 18) are oven dry weight of 2 year old crown.
Model results results represent fresh weight of 7 year old crown.
growth and yield include herbivory, disease, nutrients and
certain other cultural practices. However, due to the lack of
data, the desire to keep the model simple, and the qualitative
rather than quantitative nature of the model, we felt omission
of these factors would have little effect on the harvest strategies.
Results and Discussion
After a simulation model has been constructed, one is
confronted with the task of validation of results. This is gener
ally very difficult since most of the available data has gone into
the simulation’s construction. We were fortunate in that an
independent data set was made available for validation for
several of the intermediate model results (18).
Since the level of carbohydrates in the storage root was a
very important variable in our model, it was an essential param
eter to validate. In Fig. 3, the total carbohydrate reserves
uP
□
MODEL
*
O BSERVED
(1 9 7 7 -D IA M E T E R )
( 1 9 7 6 — PERCENT)
□_ 1
s
JULIAN DATE
4. Comparison of spear diameter (model) with percent marketable
spear (Shelton 18).
from our simulation are compared to data by Shelton (18).
Since the data were collected at the Michigan State University,
Horticulture Research Farm in 1977, 1977 East Lansing
weather information was used for the simulation. The dif
ferences in magnitude of the curves are of little concern
since the observed values are oven dry weight of 2 year old
crowns, whereas the model’s output represents fresh weight of
7 year old crowns. Rather, what should be considered is the
overall similarity in the shape of the curves. In this respect
there is close agreement between the similation and the ob
served values.
Another important consideration is the size of the spears.
Since the length of the harvested spears is limited by the range
of acceptability of the buyers4, the only other variable to
determine spear size is diameter. No data could be found on
the average spear diameter throughout a season -only on %
marketable spears. In Fig. 4, the observed % marketable spears/
harvest (18) is compared to the mean spear diameter as calcu
lated in the simulation. Here validation can only be made
through deductive reasoning. Since the % of marketable spears
refers to the % of spears >0.5 cm, it is therefore a function
of spear diameter. As this % decreases, the average spear diame
ter must also decrease. Since the curves are approximately
similar, one can conclude an adequate fit.
Weight/spear and number of spears/plant must also be
examined. The average weight/spear from the simulation, using
the 1977 East Lansing weather, varied from 35.4(± 4.97 s e )
to 41.3 (± 5.35 s e ) g/spear for the 5 plants simulated. Takatori
et al. (19) reported 44.0 (± 1.26 s e ) and 44.4 (± 1.35 s e )
g/spear from 4 year old crowns at 2 nitrogen levels. This is in
close agreement with our simulation results. From these simu
lations, the average of 25.6 (± .87 s e ) spears/plant is well
within the range reported by Ellison and Schermerhorn (5)
of 20 to 28 and 18 to 32 for 4 and 5 year old crowns, respec
tively.
To evaluate the long term effects of harvest strategies on
yield, several multi-year simulations were run using various
harvest strategies. Weather information for East Lansing, Michi
gan from 1959 to 1973 was used for these simulations. Each
strategy was replicated 5 times to represent 5 plants. These
replications will differ somewhat because each replicate has
a unique random emergence of spears which will attain harvestable size at different dates and may suffer greater or lesser
freezing losses.
Fig. 5 represents the average yearly yield of 5 plants simu
lated over this 15 year period harvested every year until June
10. This illustrates the variation in yield between years and the
range of output for the model, which is in agreement with other
authors (6 ,5 ,4 , and 11).
According to this 15 year simulation, the greatest average
yield/year is obtained by harvesting every year but terminating
harvest on June 1 (Table 2). If harvest is terminated either too
early (May 21) or too late (June 10), then the average yearly
yield is much less. With the exception of harvesting every year
until June 10, skipping strategies produced significantly lower
average yearly yield than harvesting every year (Fig. 6). Skip
ping strategies, however, did produce greater yield/plant, espe
cially when harvested past June 1 (Table 2). The cause of the
reduction of average yearly yield through skipping strategies
is generally due to the lower number of harvests (up to harvest
only 50% of the time). The increased yield/plant is due to the
resting, which allows buildup of carbohydrate reserves. Analy
sis of variance for yield/plant indicated highly significant
effects on yield due to harvest strategy (F348 = 25.35, P < 1%)
and last date of harvest (F2?48 “ 123.49, P < 1%). Highly signifi
cant effects on yield/year were also observed with harvest
strategy (F348 = 702.83, P < 1%) and last date of harvest (F248
= 481.18, P < 1%).
Through careful choice of harvesting strategies, i.e. skipping
Fig. 5. A 15 year simulation of average yearly yield of 5 asparagus
plants harvested every year until June 10 (mean ± 1 S E ).
years of terminating harvest early, a grower should be able to
maximize asparagus yield. The simulation suggests that ending
harvest on June 1 (2 or 3 weeks before buying stops) provides a
greater long term yield than allowing the plants to produce
spears until the canners terminate their purchases. One factor
that may have significant impact on the strategy used over many
years is the price the growers receive for their crop. Losing
much of a crop to severe weather (e.g. spring freezes) may
tempt the grower to continue harvesting past June 1. The eco
nomics of asparagus production could readily be incorporated
into the model by inserting alternative strategies dependent up
on data describing the economic situation of a particular year.
Table 2. Effects of various harvest strategies on mean yield per plant and
mean yield per year from 15 year simulations.
Last harvest
date
Harvest
strategy
Mean yield
per plant
(g)
Mean yield
per year
(g)
May 21 (141)
May 21 (141)
May 21 (141)
May 21 (141)
June 1 (151)
June 1 (151)
June 1 (151)
June 1 (151)
June 10 (161)
June 10 (161)
June 10 (161)
June 10(161)
Every year
1 out of 2 years
2 out of 3 years
3 out of 4 years
Every year
1 out of 2 years
2 out of 3 years
3 out of 4 years
Every year
1 out of 2 years
2 out of 3 years
3 out of 4 years
822.718 DEZ
626.184 G
858. 094 D
722.880 F
1008.834 BC
935.726 C
1098.852 A
982.382 BC
766.432 EF
1037.276 AB
1003.298 BC
817.972 DE
822.718
360.560
570.148
608.774
1008.834
582.344
754.240
781.088
766.432
677.416
713.694
642.692
H
A
B
C
1
B
G
G
G
E
F
D
zMean separation within columns by Student-Newman-Kuels, 5% level.
to
threshold and leaf weight. Finally, much historical data is
needed from long-term validation plots, monitored under dif
ferent controlled inputs.
Although the model at present is far from the level of com
pleteness that would be readily useful to the farmer, it does
serve to point towards needed research, and it suggests the
testable hypothesis that shorter harvesting periods may maxi
mize long-term yield.
Literature Cited
Fig. 6. The effects of various harvest strategies on average yearly yield
for a 15 year simulation of 5 plants. Points followed by same letter
are not indistinguishable by Student-Newman-Kuels, 5 % level.
Conclusions
The model predicts yields of spears which are quite close to
field values (16, 5, 4, and 11). However, the purpose of the out
put is not so much absolute yield prediction as harvest strategy
evaluation. A limitation of the latter is that this is a maximal
yield model, and not concerned with catastrophe-avoidance
which a farmer might wisely prefer to minimize his risk of
crop failure at the cost of returning sub-optimal yields.
More research is needed to clarify biological mechanisms
and make validation of this or any other asparagus model
possible. The only across-seasons state variable at present is
storage root carbohydrate level. This needs to be understood
as a composite function of available water and nutrients, insects,
disease and so forth, instead of merely calendar date and har
vesting strategy as at present. Storage root growth, bud site
proliferation, pre-emergent growth of spears through the harvest
season, and fern growth need more adequate formulations. More
research is needed to describe the functional relationships
among root weight, root age, and fern photosynthetic potential
as affected by light intensity, photoperiod, developmental
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