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Seasonal variation of the sprouting ability of rhizome/root buds and concentrations of storage compounds in Calystegia sepium (L.) R.Br. and Convolvulus arvensis L.

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25. Deutsche Arbeitsbesprechung über Fragen der Unkrautbiologie und –bekämpfung, 13.-15. März 2012, Braunschweig
694 Julius-Kühn-Archiv, 434, 2012
Seasonal variation of the sprouting ability of rhizome/root buds and
concentrations of storage compounds in Calystegia sepium (L.) R. Br. and
Convolvulus arvensis L.
Jahreszeitliche Veränderung der Austriebsfähigkeit der Rhizom- und Wurzelknospen und der
Speicherstoffkonzentrationen von Calystegia sepium (L.) R. Br. und Convolvulus arvensis L.
Leonie Willeke1*, Hansjörg Kraehmer3, Roland Gerhards1 & Wilhelm Claupein2
1Institute of Phytomedicine, University of Hohenheim, 70593 Stuttgart, Germany
2Institute of Crop Production and Grassland Research, University of Hohenheim, 70599 Stuttgart, Germany
3Bayer CropScience; Industrial Park, 65926 Frankfurt am Main, Germany
*Corresponding author, Leonie.Willeke@uni-hohenheim.de
DOI: 10.5073/jka.2012.434.090
Summary
Convolvulus arvensis (CONAR) and Calystegia sepium (CAGSE) are widespread perennial weeds. Our field results
demonstrate that the early application of herbicides usually does not result in complete control. Seasonal
changes of the sprouting ability of rhizomes or root buds and the transport of different organic compounds
within plants were therefore analyzed as possible indicators for times of maximum phloem and herbicide
transport. Rhizome and root pieces from fields in Southern Germany were used. CAGSE buds remained
completely dormant until March. Their sprouting ability increased up to over 90 % in May. CONAR buds
sprouted only from May onwards and reached a maximum of over 90 % in June. After September, the sprouting
ability decreased sharply until the underground organs became dormant in November.
Greenhouse experiments were carried out to analyze the carbon and nitrogen concentration of roots and
rhizomes with isotopic ratio mass spectrometer analysis (IRMS), sugars (fructose, glucose, sucrose) by using
high-performance liquid chromatography (HPLC) and starch concentrations by polarimeter. Experiments
started at the time of first shoots in spring and finished with the onset of winter dormancy in fall. Our results
show that CAGSE exhausts starch in rhizomes up to mid-June and CONAR in roots up to late June. Thereafter,
the amount of storage compounds increased again. By October, the starch concentration of dry weight in roots
and rhizomes reached 11-12 % more than in spring. The November values were 21 % and 18 %, respectively.
Sugar concentrations vary during the growing period from 1.5 % to 6 % related to dry weight. At the end of
June, an increase was recorded in parallel to starch accumulation. These results are discussed as parameters for
the control of both species.
Keywords: Bindweed, carbon, nitrogen, perennial weed, starch, sugar
Zusammenfassung
Convolvulus arvensis (CONAR) und Calystegia sepium (CAGSE) sind weit verbreitete mehrjährige Unkräuter.
Unsere Feldversuche haben gezeigt, dass eine frühe Herbizidapplikation in der Regel zu keiner vollständigen
Bekämpfung führt. Aus diesem Grund wurden die saisonalen Veränderungen der Austriebsfähigkeit von
Rhizom- und Wurzelknospen und der Transport verschiedener organischer Verbindungen in den Pflanzen als
mögliche Indikatoren für die Zeiten des maximalen Phloem- und Herbizidtransports analysiert. Es wurden
Rhizom- und Wurzelstücke von Ackerflächen in Süddeutschland verwendet. Die Rhizomknospen von CAGSE
blieben bis März dormant. Ihre Austriebsfähigkeit stieg im Mai auf über 90 %. Die Wurzelknospen von CONAR
trieben ab Mai aus und erreichten ein Maximum von über 90 % im Juni. Die Keimfähigkeit fiel ab September
stark ab, bis im November die unterirdischen Organe vollständig dormant waren.
Gewächshausversuche zur Bestimmung der Kohlenstoff- und Stickstoffkonzentration in Wurzeln und Rhizomen
wurden durchgeführt. Die Analysen erfolgten mit einem Isotopenverhältnis-Massenspektrometer (IRMS),
Zuckerkonzentrationen (Glucose, Fructose, Saccharose) wurden mittels Hochleistungs-Flüssigchromatographie
(HPLC) gemessen, Stärkekonzentrationen mit dem Polarimeter. Die Experimente begannen mit dem ersten
Austrieb im Frühjahr und endeten im Herbst zu Beginn der Winterruhe. Unsere Ergebnisse zeigen, dass die
untersuchten CAGSE-Rhizome bis Mitte Juni und CONAR-Wurzeln bis Ende Juni vermehrt Stärke abbauten. Im
Anschluss stieg die Speicherstoffkonzentration wieder an. Im Oktober lag die Stärkekonzentration bezogen auf
die Trockenmasse in den Wurzeln und Rhizomen um 11-12 % höher als im Frühjahr. Im November lagen die
Werte bei 21 % bzw. 18 %. Zuckerkonzentrationen schwankten bezogen auf das Trockengewicht während der
Vegetationsperiode zwischen 1,5 % und 6 %. Ende Juni wurde wie bereits bei der Stärke eine Zunahme
25th German Conference on Weed Biology and Weed Control, March 13-15, 2012, Braunschweig, Germany
Julius-Kühn-Archiv, 434, 2012 695
gemessen. Die Ergebnisse werden als Parameter der Bekämpfung der beiden Arten diskutiert.
Stichwörter: Kohlenstoff, perennierende Unkräuter, Stärke, Stickstoff, Winden, Zucker
1. Introduction
CAGSE and CONAR are perennial weeds, which can twine around crop plants and break down their
stems. The two species are distributed worldwide. Especially CONAR is ranked among the most
aggressive weeds for years (HOLM et al., 1977). In contrast to the small knowledge about the damage
caused by CAGSE, the economic impact of CONAR is well described. According to BOLDT (1998), the
estimated loss through bindweed in the USA amounts to over $ 377 million per year (BOLDT et al.,
1998). On sites with high weed densities, yield losses were recorded at 50-60 % for CONAR (CALLIHAN
et al., 1990). Both species can overwinter after the aboveground parts die in fall. The main and lateral
roots become woody and starch and sugar are stored (KOGAN, 1986; WIESE and PHILIPPS, 1976).
Dormant rhizome and root buds enable perennial plants in temperate climates to survive even under
adverse environmental conditions and to slow or increase the active growth in dependence of the
seasonal climatic change (MCALLISTER and HADERLIE, 1985). Without a proper control strategy, the two
species increasingly expand their root or rhizome system from year to year. Thus, a CONAR nest can
enlarge in one year on average 3 m (FRAZIER, 1943). The formation of extended patches and the high
variability of the sprouting time of rhizomes or roots in spring significantly hamper weed control.
CAGSE and CONAR can be found in both conventional and organic farming. Above all, the increasing
use of reduced tillage leads to an increase in perennial weeds (NKURUNZIA et al., 2003). In conventional
agriculture, specific herbicides therefore have to be used for targeted reduction. The above-ground
shoots can easily be killed, but only systemic herbicides can prevent the regrowth of the
underground parts (RASK and ANDREASEN, 2007).
In our field experiments, early herbicide application did not result in long-term control of the two
bindweeds. Many rhizome and root pieces of the clones formed new shoots in the same or in the next
growing season despite herbicide applications. The effective implementation of good control
strategies at the right time requires an advanced knowledge about the biology of bindweed.
Studies of NKURUNZIZA (2010) about the perennial species Cirsium arvense have already shown that
storage substances are used to form new shoots at the beginning of the vegetation period. High
photosynthesis rates seem to enhance the carbohydrate transport into rhizomes and roots.
Manufacturers, extension service and university programs recommend to apply herbicides to
actively-growing bindweeds. Some refer to the enhanced phloem transport. We assume that
systemic herbicides are more efficiently translocated into belowground organs when photosynthetic
transport is high. Based on this assumption, we examined the sprouting ability of the rhizome and
root pieces on the one hand and the concentration variation of stored nutrients like sugars and starch
on the other hand as well as the carbon and nitrogen concentration during one growing season.
2. Materials and methods
2.1 Analysis of the sprouting ability of rhizome and root pieces
From March 2010 until the dormancy of rhizome and root pieces in the end of November, pieces with
six nodes and a length of about 10 cm were collected each month. The bindweed originated from a
field on the Ihinger Hof (Experimental Station of the University of Hohenheim, Germany) and hedge
bindweed from a field near Uhingen (Germany). After collection, the underground organs were
washed, wrapped in filter paper and placed in a pot with water. The pots were placed in a greenhouse
with a day-night cycle of 12 h / 12 h at 23 °C. After seven days, the number of sprouted pieces was
noted and documented as a percentage of totally assessed plants.
2.2 Analysis of different ingredients of rhizome and root pieces
At the beginning of the vegetation period (April 2010), vegetatively propagated root and rhizome
pieces with 6-7 nodes of field bindweed and hedge bindweed were planted separately in 13 x 13 x
25. Deutsche Arbeitsbesprechung über Fragen der Unkrautbiologie und –bekämpfung, 13.-15. März 2012, Braunschweig
696 Julius-Kühn-Archiv, 434, 2012
22 cm pots. The plants were grown in a clay-substrate mixture (pH 7.2) and placed in the outside area
of a vegetation hall of the University of Hohenheim (Stuttgart, Germany). The growth conditions such
as day-night cycle and temperature corresponded to ambient conditions. The plants were
additionally watered when necessary. On the first and fifteenth of each month, six plants were
collected, the roots freed from soil and the concentration of the ingredients analyzed as described
below.
Carbon and nitrogen content were measured by using isotope ratio mass spectrometry (IRMS). For
this analysis, the root samples were dried for three hours at 80 °C and mortared in liquid nitrogen to a
fine powder. Samples of 2-3 mg were weighed into tin capsules (5 x 9 mm). IRMS was performed on a
Thermo Finnigan Delta plus XP system, coupled to a Euro EA elemental analyzer (Euro Vector
Instruments and Software, Hekatech, Wegberg, USA) (oxidation furnace, 1000 °C, reduction furnace,
650 °C, carrier gas, 40 kPa; packed column temperature, 90 °C). As standard material, acetanilides
were utilized. For data acquisition and processing, Thermo Electron ISO Date NT software, version 2.0,
was used.
In preparation of the samples for the polarimetric starch determination by Baumann Grossfeld, 2.5 g
of fresh rhizome and root pieces were homogenized with an Ultra-Turrax, hydrochloric acid was
added and heated in a water bath at 100 °C. Then a Carrez clarification was conducted to precipitate
unwanted substances such as proteins. Finally, the samples were filtered and measured on a Perkin
Elmer polarimeter.
For the high-performance liquid chromatography (HPLC) measurement of sugar analysis, 2.5 g of the
rhizome and root pieces were first homogenized in a water-acetonitril mixture by using an Ultra-
Turrax, then centrifuged and the pellet was discarded. The analysis was performed on a Perkin Elmer
HPLC system (evaporative light scattering detector (ELSD), using a HPLC column (ShodexAsahipak
NH2P-50, 250 x 4.6 mm, 5 microns), flow rate 1 ml/min; demineralized eluent, acetonitrile and distilled
water).
2.3 Statistical analysis
The experiments were conducted in a completely randomized block design. Analysis of variance
(ANOVA) was performed on the data from both experiments. The statistical analysis was conducted
using SAS 9.2. and the graphics were created with Excel 2007. For the analysis of the root ingredients
we used the glm procedure for pairwise comparisons to test for significant variations of different
months in vegetation periods on the response variables of carbon, nitrogen or carbohydrates.
3. Results
3.1 Sprouting ability of rhizome and root buds
Figure 1 shows the sprouting ability of the rhizome and root pieces of the two perennial species. The
onset of sprouting ability was recorded in April. At this month, the rhizome sprouting ability of CAGSE
was 15 % and reached a maximum of 98 % in May. In contrast to the rhizomes, the roots were
dormant until May. In June, a large number of pieces (94 %) formed new shoots. The underground
organs of both weed species could sprout to a rate of over 78 % throughout the summer until late
September. In general, the sprouting ability of the 10 cm long underground organs of field bindweed
was on average 10 % higher than that of hedge bindweed. In September, the high sprouting ability
decreased significantly and CAGSE pieces became dormant in November and CONAR pieces in
December.
2
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25. Deutsche Arbeitsbesprechung über Fragen der Unkrautbiologie und –bekämpfung, 13.-15. März 2012, Braunschweig
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at the flowering period, the lowest concentration of less than 1 % was measured. From the end of
flowering, in mid-August, the nitrogen content increased again. By the end of the growing season, it
changed only slightly and the contents no longer differed significantly.
Fig. 3 Nitrogen content (% dry weight) in the rhizomes of CAGSE (left panel) and roots of CONAR (right
panel) from early May until mid-November. Values indicated by different letters are significantly
different (P < 0.05).
Abb. 3 Stickstoffkonzentration (% Trockengewicht) der CAGSE-Rhizome (linke Graphik) und CONAR-Wurzeln
(rechte Graphik) von Anfang Mai bis Mitte November. Mit unterschiedlichen Buchstaben gekennzeichnete
Balken unterscheiden sich signifikant (P < 0,05).
3.5 Analysis of starch content
Figure 4 shows that, in both species, the starch concentration in the rhizomes and roots was highest
at the beginning of planting time compared to the other months. The starch content of CONAR roots
increased slowly to a minimum of 12 % at 15.06. The CAGSE rhizomes possessed the lowest starch
concentrations of 9-13 % by mid-July. These differed significantly from the levels measured at other
times.
An increased storage of starch occured in both species at early to mid-July. From the end of the
flowering period, the content varied only slightly and no longer changed significantly. In general, the
figure shows that CAGSE as well as CONAR stored about 20 % starch as a reserve.
Fig. 4 Starch content (% dry weight) in the rhizomes of CAGSE (left panel) and roots of CONAR (right panel)
from early May until mid-November. Values indicated by different letters are significantly different
(P < 0.05).
Abb. 4 Stärkekonzentration (% Trockengewicht) der CAGSE-Rhizome (linke Graphik) und CONAR-Wurzeln (rechte
Graphik) von Anfang Mai bis Mitte November. Mit unterschiedlichen Buchstaben gekennzeichnete Balken
unterscheiden sich signifikant (P < 0,05).
25th German Conference on Weed Biology and Weed Control, March 13-15, 2012, Braunschweig, Germany
Julius-Kühn-Archiv, 434, 2012 699
3.5 Analysis of sugar content
The sugar concentrations of both species strongly changed during the vegetation period. In the early
stages of growth, especially the glucose level raised in the rhizome and root pieces from
approximately 0.5 and 0.8 % to 4 % and 2 %. Figure 5 illustrates that at the time of flowering and seed
production the total sugar concentration of the underground organs of both weeds decreased
significantly. In mid-August, the lowest concentrations of approximately 1.5 % were measured. From
September onwards, sugar was increasingly stored - at this time, in both species, almost exclusively in
the form of sucrose.
Fig. 5 Sugar content (% dry weight) in the rhizomes of CAGSE (left panel) and roots of CONAR (right panel)
from early May until mid-November. Values indicated by different letters are significantly different
(P < 0.05).
Abb. 5 Zuckerkonzentration (% Trockengewicht) der CAGSE-Rhizome (linke Graphik) und CONAR-Wurzeln (rechte
Graphik) von Anfang Mai bis Mitte November. Mit unterschiedlichen Buchstaben gekennzeichnete Balken
unterscheiden sich signifikant (P < 0,05).
4. Discussion
The two studies show the percentage of sprouting ability of rhizomes and root pieces on the one
hand and the trend of ingredient concentrations of the underground storage organs of the two
species CAGSE and CONAR on the other hand. Although in the first experiment the root and rhizome
pieces were collected from the same nests (which could suggest a possible cloning), there was a high
variability in the sprouting ability in the respective months. The sprouting ability was highest from
May (CAGSE) and June (CONAR) to August. But never did 100 % of the rhizomes or roots sprout. This
high variability of sprouting time during the growing season seems to be a major survival strategy of
the plants after herbicide application. This applies especially to early applications before the buds of
the rhizome and root pieces sprout and thus the herbicide cannot adequately control the
underground organs. In addition to the inhomogeneous sprouting in the second experiment, an
inhomogeneous shoot growth of the different plants of one replication was found.
At the beginning of the growth period the stored nutrients were mobilized. The starch granules were
presumably reduced and transported as an energy source into the developing stem (sink). The
resulting decrease of the starch in underground parts until mid-June is illustrated in Figure 5. Only
through increased photosynthesis with increasing shoot growth in June, the source and the sink
might have reversed after reaching their compensation point. Assimilates were now transported into
the roots and stored in form of sugars and starch. The described transport directions of starch are
indicated by measured sugar concentrations. As Figure 4 shows, the glucose level rised sharply,
especially in CAGSE rhizomes from mid-June to early July. This could be due to the fact that starch
cannot be transported through the phloem, the plants have to convert the starch with the help of
phosphorylases and amylases into glucose. In addition, from August onwards, the sucrose
concentration increased sharply in the underground parts of both weeds. This is consistent with the
theory that the transport of starch in other parts of plants takes place mostly in the form of sucrose
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(HELDT, 2003).
The measured carbohydrates and the total carbon and nitrogen analyses underline the seasonal
trend of source and sink relationships. In addition to hydrogen and oxygen, carbon is one of the main
components of organic compounds in plants (CAMPBELL, 1997) and is very versatile. At the beginning
of the growing season, multiple carbon compounds are required as an energy source for growth.
Thus the concentration declined first, as can be seen in Figure 2. Presumably only through growth-
increased photosynthetic activity with CO2 fixation in combination with basipetally increased
transport of various carbon compounds, did the carbon concentration increase again in August. Also,
nitrogen is an important component in plants. It occurs mainly in nucleic acids, proteins, hormones
and coenzymes (CAMPBELL, 1997). Proteins also serve alongside carbohydrates and fats as an energy
source and serve as fuels of respiration. This could be a reason for the measured decrease in nitrogen
content in the underground organs during the shoot growth and the increase only by mid-August
(Fig. 3).
Studies that expand knowledge about the biology of perennial species and especially on the sources
and sinks of carbohydrate dynamics promise to improve weed management strategies.
Compensation point is a pivotal time because it determines the start of increased weed-crop
competition (NKURUNZIA, 2010). The right time of herbicide application is very important. Growth
stages of the plants, soil moisture and precipitation play an important role (WESTRA, 1992). In recent
years, various authors have given recommendations for the optimal time of herbicide treatment.
Several studies indicate that herbicide treatments for herbicides such as glyphosate and 2,4-D should
take place during the first flowering, when carbohydrate concentration is low (ALCOCK and DICKINSON,
1974; CALLIHAN et al., 1990; KOGAN, 1986; PETERSON, 1998). However, carbohydrate concentrations were
not measured in these experiments. It may well be that by observing the actual trend of reserve
concentrations, application timing can be optimized. From our studies we can conclude that the
compensation point occurs probably before flowering. The starch concentration has reached its
minimum 1.5 months before flowering begins. Thus it can be assumed that optimal treatment should
take place shortly after the compensation point when the plant has little reserves and when the
translocation of the herbicides through the phloem with assimilates into the underground organs can
be expected. In addition to the application of herbicides, repeated tillage during the growing season
can reduce weed infestation. The formation of new shoots after harrowing cuts depends on the
reserves (HAKANSSON, 2003). Thus, in this case, the compensation point is an important time for
mechanical control. The reserves of rhizomes and roots can quickly be removed with this strategy.
It should be noted, however, that data from field trials can differ significantly in comparison to
greenhouse experiments. Unlike annual weeds, which are grown from seed and thus demonstrate a
clearly defined development, perennial species often show large discrepancies as a result of nest-
building with extensive underground systems and high phenotypic variability. Therefore, it is
necessary to perform further field studies under different environmental conditions. To optimize
control strategies, it is important to find out the influence of biotic and abiotic factors on the
dormancy of rhizomes and roots, which also affect the seasonal trend of source-sink relations of the
reserves.
Acknowledgments
Wewouldliketothank F. Walter, B. Hoeglinger (Institute of Phytomedicine, University of
Hohenheim), W. Armbruster and A. Lang (Institute of Food Chemistry, University of Hohenheim) for
their technical assistance during analysis of the storage compounds. This study was supported by
Bayer CropScience.
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HELDT, H. W., 2003: PFLANZENBIOCHEMIE., SPEKTRUM AKADEMISCHER VERLAG HEIDELBERG, BERLIN.
NKURUNZIA, L., E. ROSEQVIST AND J.C. STREIBIG, 2010: PHOTOSYNTHESIS AND GROWTH OF NEWLY ESTABLISHED SHOOTS OF CIRSIUM
ARVENSE AND TUSSILAGO FARFARA ARE RESOURCE INDEPENDENT. WEED RESEARCH 51, 33–40.
NKURUNZIZA, L., 2010: PHENOLOGY AND SOURCE-SINK DYNAMICS OF CARBOHYDRATES IN RELATION TO MANAGEMENT OF PERENNIAL
WEEDS: CIRSIUM ARVENSE (L.) SCOP AND TUSSILAGO FARFARA L., PHD THESIS, DEPARTMENT OF AGRICULTURE AND ECOLOGY,
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PETERSON, D. AND P.W, STAHLMANN 1989: FIELD BINDWEED CONTROL IN FIELD CROPS AND FALLOW. KANSAS STATE UNIVERSITY
AGRICULTURAL EXPERIMENT STATION AND COOPERATIVE EXTENSION SERVICE, 1-12.
RASK, A. M. AND C. ANDREASEN, 2007: INflUENCE OF MECHANICAL RHIZOME CUTTING, RHIZOME DRYING AND BURIAL AT DIFFERENT
DEVELOPMENTAL STAGES ON THE REGROWTH OF CALYSTEGIA SEPIUM. WEED RESEARCH 47, 84–93.
WESTRA, P., P. CHAPMAN, P.W. STAHLMAN, S.D. MILLER AND P.K. FAY, 1992: FIELD BINDWEED (CONVOLVULUS ARVENSIS) CONTROL
WITH VARIOUS HERBICIDE COMBINATIONS. WEED TECHNOLOGY 6, 949-955.
WIESE, A. F. AND W.M. PHILLIPS, 1976: FIELD BINDWEED. WEEDS TODAY 6, 22–23.
... As great bindweed is deciduous, it is assumed that materials from most leaves are translocated down to the rhizome system in late autumn. In spring and summer, leaves in upper parts of the stem send most sugars towards the shoot tip as part of the sink source relationship (Willeke et al. 2012). Thus it was expected that herbicide applied to these leaves in late autumn would move to the rhizomes as effectively as herbicide applied to lower leaves, which was conirmed with the present work. ...
... Results from the ield trial suggest that it would not be wise to wait until the following autumn to respray the regrowth as plants could recover fully by then. However, respraying as soon as shoots appear may not be effective as sugar low would be mainly upwards from rhizomes to new shoots initially (Willeke et al. 2012). It may be better to wait until translocation of sugars back down to the rhizomes has begun again so herbicides might be moved into the rhizomes, although only basal leaves would be expected to send herbicides to the rhizomes at this stage. ...
Article
Full-text available
A ield trial was conducted in Palmerston North to compare autumn applications of several translocated herbicides for great bindweed (Calystegia silvatica) control in riparian zones. Regrowth in the following spring showed that a triclopyr/picloram/ aminopyralid mixture, a 2,4-D/dicamba mixture and aminopyralid by itself were the three most effective treatments, though none gave complete control. Glyphosate provided partial control whereas metsulfuron and clopyralid provided poor control. These and two other herbicides were further assessed in a glasshouse trial in which they were applied to leaves either on the upper or lower half of plants to compare eficacy. The relative effectiveness of these herbicides on great bindweed was similar to that found in the ield. Most herbicides had similar eficacy whether applied to upper or lower parts in autumn, except glyphosate, which was more effective applied to upper plant parts. Implications for control of great bindweed in riparian plantings are discussed.
... Another trend is to focus more on perennial weeds' vegetative reproduction regard ing dormancy and sprouting in weed control studies in Europe and especially in Scandinavia (Brandsaeter et al., 2010;Willeke et al., 2012;Andersson et al., 2013). ...
Chapter
Seed biology is important for emergence in the field and for future weed infestations. This chapter focuses on seed biology, germination, dormancy and efforts in predicting weed emergence from seeds from a European perspective. It presents a brief overview of population dynamics in time and space, the factors influencing the dynamics and how population dynamics can be modelled. Emergence from the seed-bank starts with germination, pre-emergence growth and finally emergence. In addition to seeds, vegetatively propagated material is briefly mentioned. Dormancy influences under what conditions that germination can occur and regulates timing of germination. Population dynamics are important for understanding the whole system and are often based on the life-cycle of weeds: seed-bank, seedlings, adult plants, seed production and dispersal. Challenges in modelling emergence and population dynamics are large, due to differences between and within populations of species, variability in species response and there being many weed species in the same field with contrasting characteristics.
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RASK, A. M. AND C. ANDREASEN, 2007: INflUENCE OF MECHANICAL RHIZOME CUTTING, RHIZOME DRYING AND BURIAL AT DIFFERENT DEVELOPMENTAL STAGES ON THE REGROWTH OF CALYSTEGIA SEPIUM. WEED RESEARCH 47, 84–93.
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Boldt, P. E., S.S. Rosenthal AND R. Srinivasan, 1998: DISTRIBUTION OF FIELD BINDWEED AND HEDGE BINDWEED IN THE USA. JOURNAL OF PRODUCTION AGRICULTURE 11, 377-381.
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MCALLISTER, R.S. AND L.C. HADERLIE, 1985: SEASONAL VARIATIONS IN CANADA THISTLE (CIRSIUM ARVENSE) ROOT BUD GROWTH AND ROOT CARBOHYDRATE RESERVES. SCIENCE 33, 44-49.
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