沙漠研究 22-1, 13 -16 (2012 )"ICAL 1 / DT X"

Journal of Arid Land Studies－Refereed Paper－

Root System Development of Prosopis Seedlings
under Different Soil Moisture Conditions
Kiyotsugu YODA*1), Mohamed Abd ELBASIT2), Buho HOSHINO3),
Hiroshi NAWATA4) and Hiroshi YASUDA2)
Abstract: Prosopis juliflora (Swartz) DC is one of the most economically and/or ecologically important legumes in arid andsemi-arid regions of the world. Prosopis was introduced into Sudan in early 20th century, and succeeded in controlling the expansionof desert in northern Sudan. Recently, however, this species distributes widely and rapidly in eastern Sudan, and induces many
problems in human subsistence. To clarify the mechanism of rapid invasion of Prosopis, we investigated the response of seedlingroots to different soil moisture conditions. Prosopis seedlings were cultivated twice (seven weeks each) in Desertification
Mechanism Analysis System of Arid Land Research Center, Tottori University. Seedlings were cultivated in columns with sandy
soils and dripping irrigation once per day. Second experiment was done in columns with four drip irrigations (top, top and middle,middle and bottom, or bottom of columns). Seedlings were carefully harvested, and dry weight and root structure were analyzed.
Results were (1) seedlings developed two root systems, one was horizontal root just below the soil surface, and the other was tap root
penetrated beyond 1m depth, (2) water supply on the surface increased dry weight and total root length of seedling, (3) seedlingdeveloped lateral roots vigorously at higher soil moisture areas. These results suggest that seedling responds sensitively to theincrease of soil moisture after the rain, and develops root system flexibly against patches of soil moisture. These properties may
enable Prosopis to invade easily into new habitats and distribute widely and rapidly in eastern Sudan.
Key Words: Prosopis juliflora, Root, Soil moisture, Sudan

1. Introduction
Prosopis juliflora (Swartz) DC is an evergreen legumenative to South to Central America and the Caribbean, and oneof the most economically and/or ecologically important trees inarid and semi-arid regions of the world. This species wasintroduced into North-east Africa in early 20th century(Pasiecznik et al., 2001) to arrest the desertification andimprove agricultural schemes. In northern Sudan, theexpansion of desert has been successfully controlled bypreparing the shelterbelt of Prosopis (Laxén, 2007). Recently,
however, this species distributes widely and rapidly in eastern
Sudan, and induces many problems for human subsistence inthis country as follows; it crowds out native and edible plants,
blocks drains and irrigation canals, and is a major menace forfarmers; its thorns are harmful both to farm workers and theirmachinery; it consumes underground water; its pods bringabout some animal diseases (Mwangi and Swallow, 2005;
UNEP, 2007).
Some properties relating the invasion of Prosopis are as
follows: 1. Tree has low branching and highly coppicing
abilities (Bosu et al., 2009), 2. Trees flower at 3-5 years ofage generally, and under certain experimental condition, begin
flowering at 3-4 months after germination (Pasiecznik et al.,2006), 3. Trees develop two distinct root systems as surface
lateral roots and deep tap roots (Jenkins et al. 1987, Pasieczniket al., 2001), which may have a role of hydraulic redistribution(Hultine et al., 2004), and cause of survival in drier condition
without obvious xeromorphic adaptations (Bristow, 1996), 4.
allelopathic effects of leaves inhibit the growth of other plants(Nakano, 2010), 5. seeds are rapidly and easily dispersed bylivelihoods and other animals (Berhanu and Tesfaye, 2006),
and so on. All or parts of them might facilitate the invasion of
Prosopis into eastern Sudan, and the establishment ofappropriate management plans of Prosopis is keenlydemanded.
We investigated the growth pattern of Prosopis seedlings indifferent irrigation conditions. The purpose of this study is toclarify the factor of Prosopis invasion, focusing on thearchitecture and development of roots.

2. Materials and Methods
This study was done in Desertification Mechanism
Analysis System (DMAS) of Arid Land Dome, Arid Land

* Corresponding Author. yoda@isenshu-u.ac.jp
Minamisakai Shinmito 1, Ishinomaki, Miyagi 986-8580, Japan
1) Faculty of Science and Engineering, Ishinomaki Senshu University

3) Faculty of Environment System, Rakuno Gakuen University2) Arid-Land Research Center, Tottori University

4) Research Institute for Humanity and Nature

Research Center, Tottori University. Environmental conditionsin DMAS were set as follows: air temperature, 27˚C; airrelative humidity, 40%; wind speed, 0.85 m/sec.
Twelve columns (diameter 45cm, depth 1m) of polyvinyl
chloride were prepared in DMAS and filled with sandy soil.
Prosopis juliflora seeds were disseminated on 27 April 2010,germinated and nursed in an incubator beforehand. Seedlings
with four mature leaves were transplanted in the columns (oneseedling per column) on 21 May 2010, and cultivated with drip
irrigation (5 minutes, once per day). Seven seedlings wereharvested on 7 July 2010. Sandy soils around roots of theseedlings were carefully removed under running water, andlengths of shoots were measured. To examine the distributionof roots, tap roots were rearranged vertically, divided intosegments of 20cm-intervals from the soil surface downward,
and the lengths of lateral roots attached to each segment weremeasured. All these parts were oven-dried (60˚C, 24 hours),
then weighed with electric balance.
As a second experiment to investigate the root
developments in different soil moisture conditions, otherpolyvinyl chloride columns (diameter 15 cm, depth 1.3 m)
were prepared in DMAS. Four types of irrigation conditions
were set up as follows: A, top irrigation in the column; B, top
and middle (depth 0.6 m) irrigations; C, middle and bottom(depth 1.2 m) irrigations; D, bottom irrigation. Threecolumns were prepared for each irrigation treatment (total 12columns). Prearranged seedlings, disseminated on 4 August
2010, were transplanted in the columns on 10 September 2010,
cultivated with drip irrigation (10 minutes, once per two days),
harvested on 27 October 2010, and measured in the samemanner to the previous experiment.
Differences of dry weight or root length among theirrigation treatments were tested by Tukey-Welsch stepwise
multiple comparison.

3. Results and Discussion
3.1. Root weight and length distribution
In the first experiment, seedlings expanded 11~13 leaves inseven weeks after transplantation. Some seedlings penetrated
single tap root beyond 1 m. Others formed two- or threebranched tap root, maximum total length of them reached 2.3
m. Averaged shoot length of seven seedlings was 19.0 ± 3.7
cm. Mean value of total root length of them was 184.1 ± 35.0
cm, and 9.8-times longer than that of shoot (Table 1).
Averaged dry weight of shoots was 0.34 ± 0.092 g, whereasthat of roots was 0.15 ± 0.035 g, which was a half of shootweight. All seedlings expanded lateral roots vigorously justbelow the soil surface (Fig. 1). Some seedlings formed lateralroots also at the bottom of column, because of moistened

Table 1. Length, weight and Root/Shoot ratio of Prosopis seedlings(Average ± S.D).
Length (cm)
Root
Shoot
19.0 ± 3.7

184.1 ± 35.0

Weight (g)
Shoot
Root
0.34 ± 0.092

0.15 ± 0.035

Root/Shoot ratio
LengthWeight
9.8 ± 1.6

0.45 ± 0.10

Fig. 1. Root length distributions along 20 cm-interval in thickcolumns.
Bars with different letters indicate significant
differences between data sets (n = 7, P < 0.05). “#” means to be no
significant difference because of large variation of data values.

condition of bottom sand.
Lateral root growth was
suppressed between 20 and 80 cm depths in the column.
Prosopis is typical phreatophyte, and has distinct two rootsystems as tap root and lateral root (Gallaber and Merlin, 2010).
We also confirmed this property in cultivated seedlings. They
formed roots about ten-times longer than shoots within sevenweeks after germination. In addition, roots were extremely
longer than that reported by Pasiecznik et al. (2001), in whichroot length of Prosopis seedling reached 40 cm in eight weeksafter germination. Vigorous root elongation observed in ourstudy ensure that Prosopis seedling responds to soil wetting
just after the rain, and penetrates tap root rapidly, which would
make possible to survive in the following dry seasons.
Glendening and Paulsen (1955) reported that nine-month-old
seedling of velvet mesquite in the field had root/shoot ratio in
dry weight as 1.99/1, whereas the ratio of ten-month-old
seedling under cultivation was 0.87/1. In our study,
two-month-old seedlings showed root/shoot ratio as 0.45/1,
and suggested that the root development under cultivation
should be suppressed. Further improvement of experimental
conditions might reveal some potential growth ability of
Prosopis root system.
3.2. Response of root system to different soil moisture
conditions.
Second experiment revealed that dry weights of seedlings
were clearly different among four irrigation treatments (Fig. 2).
Treatment “B” induced seedling to grow the most vigorously,
followed by treatment “A”. Seedling growths in treatments

Fig. 2. Total dry weight of seedlings under four irrigation treatments.
Bars with different letters indicate significant differences between
data sets (n = 3 in each treatment, P < 0.05).

Fig. 3. Total root length distributions along 20 cm- interval in depth
below the soil surface of columns under four irrigation
treatments. Bars with different letters indicate significant
differences between data sets (n = 3 in each treatment, P < 0.05).
One sampling in “D” elongated a tap root beyond 1 m (opencolumn).

“C” and “D” were not different significantly, and both thesetreatments suppressed seedling growth than the former twotreatments. Two seedlings in “A” and all seedlings in “B”
penetrated tap roots beyond 1m, whereas the length of tap rootsin “C” did not exceed 0.8 m. Tap root length in “D” variedamong three seedlings as 0.2 m, 0.6 m or over 1 m.
Root distributions in columns were also different among
the four irrigation treatments (Fig. 3). In treatment “A”,
seedlings formed lateral roots vigorously just below the soil
surface, and suppressed root development in the deeper area ofcolumns below 20 cm depth. Seedlings in treatment “B”
responded to the two irrigations (top and middle portions ofcolumns) with active lateral root developments. Difference ofroot length was statistically not significant among depths
because of various responses of three seedlings. Treatments“C” and “D” constrained root growth more than the former
two treatments.

Prosopis flexuosa modified root architecture plasticallyaccording to different soil moisture conditions, and acquired
water and nutrients in the soil effectively using horizontal
surface roots and deep tap roots (Guevara et al., 2010). P.
juliflora in our study showed the similar response, watersupply to the soil surface in treatments “A” and “B” promoted
the growth of surface lateral roots, and increased dry weightsof seedlings. In addition, the response of seedlings intreatment “B” suggested that P. juliflora has an ability tomodify root architecture in heterogeneous soil water
distribution, and acquire available water resources effectively
in the soil.
In Sudan, there are distinct dry and wet seasons in annual,
the latter continuing three months from July to September(Cabmerlin, 1997; Elagib and Mansell, 2000). Results in thisstudy show that, during the three months of the rainy season,
Prosopis juliflora seedling has potentials to penetrate tap roots
beyond 3 m depth in the soil and to grow vigorously withexpanding lateral surface roots. With acquiring theseproperties, Prosopis seedling responds to desirable watercondition as rain fall immediately, and constructs root system
rapidly prior to dry-up of soil in the following dry season. If
the root system would reach to some water-plentiful areas in
the soil as sub-aquifer during the rainy season, the seedling
could secure its establishment in new habitats even within thedry season. As a result, P. juliflora might to survive and/or
tolerate under drier environmental conditions, and invade andexpands in various areas rapidly. All these properties
detected in the earlier developmental stages of P. juliflora
seedlings would be advantageous to invade into the newhabitats and expand distribution rapidly in arid and semi-arid
environment as Sudan.
Acknowledgements
This study was carried out under the cooperative researchwith Research Project “A Study of Human Subsistence
Ecosystems in Arab societies: to Combat Livelihood
Degradation for the Post-oil Era” (Project Leader: Hiroshi
Nawata, Research Period: 2008-2013), Research Institute for
Humanity and Nature (RIHN), Japan, and the Joint Research
Program of Arid Land Research Center, Tottori University.

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