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Influence of the aqueous extracts of Ulva lactuca and Chlorella kessleri on


growth and yield of Vicia faba

Article  in  Algological Studies · July 2005


DOI: 10.1127/1864-1318/2005/0116-0213

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Algologica! Studies116 Stuttgart,July 2005

[nfluence of the aqueousextractsof Ulva lactuca and


Chlorella kessleri on growth and yield of Viciafaba

By AMAL H. EL-NAGGAR, MOHAMED E.H. OSMAN,


MOSTAFAM. EL-SHEEKH and SALY F. GHEDA

TantaUniversity,Faculty of Science,Botany Department,Tanta,Egypt

With 6 figures and 3 tables in the text

Abstract: The aqueous extracts of Ulva lactuca and Chlorella kessleri increased the per-
::entage of gennination, all seedling growth pararneters, leaf area, pigments content and
the fresh and dry weights of roots and shoots of Vivia faba compared to the controls. The
two algal extracts also increased concentrations of inorganic elements and bound nitrogen
::ontent (soluble and insoluble) in harvested ~ faba. Whereas C. kessleri extract was more
stimulatory for the accumulation of Na+ and K + in roots and shoots. U. lactuca extract in-
creased Mg2+, Ca2+ and p5+ more than did C. kessleri extract The extract of U. lactuca
was superior to C. kessleri extract in stimulating the accumulation of soluble nitrogen in
roots and of insoluble nitrogen in roots and shoots. Both of the extracts produced a higher
accumulation of insoluble nitrogen in shoots than in roots and gave higher levels of pro-
teins and carbohydrates in the pods. However, C. kessleri extract was more effective in the
accumulation of directly reducing sugars, while U. lactuca extract rather raised the
arnounts of total reducing sugars.At all concentrations tested, both algal extracts, specially
U. lactuca extract, increased yield pararneters of ~ faba, including weight and number of
fruits and of seeds. Quantitative analysis of plant hormones revealed that U. lactuca con-
tained more cytokinin and betaines than C. kessleri, -while the latter was richer in auxin
andgibberillin.

Key words: Alga! extract,Viciafaba, honnones,seedlinggrowth,yield.

Introduction
Aqueous seaweed extracts have been employed extensively as fertilizer additives
for crop plants. Their beneficial effects include the enhancementof seedgermina-
tion (AITKEN & SENN1965) by overcoming germination inhibitors (HIRSCHet aJ.
1989), stimulation of plant growth (TAMILSELVAN & KANNAN1994), reduction of
storage 10ssesof fruits (BLUNDEN1972), increased uptake of inorganic nutrients
from the soil (BLUNDEN& WOODS1969), and increased plant resistance to pests,
frost, fungal and insect attack (ARA et aJ. 1996 and Wu et aJ. 1997).

0342-1120/04/0157-211
$ 4.25
<9 2005 E. Schweizerbart'sche Verlagsbuchhandlung, D- 70176 Stuttgart
=
AlgologicaI Studies 116 Arch. Hydrobiol. Suppl. 157
214 A. H. EL-NAGGAR et al

Seaweedextract of Ascophyllum nodosum as foliar spray increased the growth


of spinach (YVIN 1994), Capsicum annuum (ERISet a!.1995) and Phaseolus luna-
tus (REnz & TRUMBLE1996). These effects appear to be due to plant growth reg-
ulators, betaines and oligosaccharides present in the extracts which regulate the
endogenous polyamine synthesis (KLOAREGet al. 1996). However, MÓLLER&
SMITH(1998) attributed the promotive effect of Ascophyllum nodosum or Lami-
naria hyperborea suspensionson the seedlings of lettuce solely to their high min-
eral content especia!ly potassium.
Seaweed biofertilizer significantly stimulated crop yield of potato (LOPEZ
MOSQUERA & PAZOS1997 and KOWALSKIet aJ. 1999). The extract of Enteromor-
pha increased germination, seedling elongation, and catalase activity in the en-
dosperm and embryo of rice seeds(MEHTAet aJ. 1999). Furthermore, EL-MELEI-
GY (1999) found lhal air-dried seaweeds (Laurencia papillosa, Acanthophora
magadiformis, Ulva lactuca and Sargassum dentifolium) increase the total bio-
mass, yield components, photosynthetic pigments and growth promoting sub-
stances of Vicia faba L. Crude extracts from green seaweeds (Cladophora dal-
matica, Enteromorpha intestinalis, Ulva lactuca) and the led algae (Corallina
mediterranea, Jania rubens, Pterocladia pinnate) were found to increase seed
germination, seedling growth, protein, tota1 soluble sugars and chlorophyll con-
tenis in Viciafaba leaves (EL-SHEEKH& EL-SAlED2000).
Few references on the effect of freshwater algal extracts on plant growth are
available. However, extracts of Chlorella vulgaris was shown to increase the per-
centage of germination and seedling growth of cotton seeds (BIL'MES& SHOTOK
1988), promoted radish growth and showed a promotive effect similar to that of
kinetin (NAKAO et al. 1994). Moreover, acceleration of seed germination and
plant growth were reported using filtrate of Phormidiumfoveolarum (WANGet aJ.
1991), Aulosirafertilissima, Spirulina subsalsa (ZEENATet aJ. 1994) and Micro-
coleus vaginatus (KHAN et aJ. 1998 and ZAKAULLAH1999).
The present work is intended to (1) compare the effect of aqueous extracts of
Ulva lactuca (green seaweed) and Chlorella kessleri (freshwater a!ga) applied
with seed soaking on the growth and yield of Vicia faba plant, (2) to eva!uate the
role of a!ga! extracts as plant growth regulators and supporting the evidence for
the occurrence of plant hormones in a!gal extracts, (3) the isolation and identifi-
cation of plant hormones in the two a!ga! extracts.

Materials and methods


The green seaweed Ulva lactuca was collected from the Mediterranean Sea beaches at
Alexandria city. Samples were kept in air-tight plastic bags, transported immediately to the
laboratory, washed several times with filtered sea water to remove epiphytes and sand par-
ticles, washed with distilled water to remove adhering solutes, dried in an oven to constant
weight at 40°C and grinded thereafter.
The fresh water alga Chlorella kessleri (strain 211-11h, Samm1ungvon Algenkulturen,
Pflanzenphysiologisches Institut, Universitat Gottingen) was cultivated in KUHL medium
(KUHL 1962). The purified Chlorella kessleri was cultured in 21 Erlenmeyer flasks, incu-
Influence Dr Ulva lactuca and Chlorella kessleri on Vicia faba 215

bated under continuous fluorescent light, aerated with air mixed with 3 % CO2. Green cells
were harvested during exponential growth phase by centrifugation at 3000 rpm for 15 min-
utes. The cell pellet was rinsed three times and resuspended in sterilized distilled water to
remove traces of growth medium (ROGERS& BURNS1994). The final biomass was oven
dried to constant weight at 40 °C, then powdered by mortar.
The aqueous algal extracts were prepared using distilled water according to the meth-
ods cited by BRAIN et aJ.(1973), BLUNDENet aJ.(1979). The concentrations used were 0.4,
0.6,0.8, 1, 1.2, and 1.4%.
Viciafaba seeds (Giza-461) were obtained from the Main Crops Improvement Station
(MCIS), Kafr EI-Sheikh. Seeds were surface-sterilized in 3.5% sodium hypochlorit~
20 minutes, rinsed several times with distilled water, then soaked for 24 hours in aqueous
extracts. Some seedswere soaked in distilled water as control. The soaked seedswere ger-
minated in sterilized Petri-dishes containing clay soil then kept at 22 °C u~er 12 h
light/dark cycle and divided into 2 groups. In ODegroup, the number of emerged seedlings
was recorded and their morphological criteria, including root length, lateral root number,
and shoot length were measured in the 2nd,4th and 8th days.
After 8-10 days, seedlings were removed carefully from the growth medium and
washed thoroughly. Leaf areas of these removed seedlings were measured using the
Ushikata x-plan 360d planimeter (FEATONBY-SMITH & VAN STADEN1983), then separated
into roots and shoots and the fresh and dry weights were recorded. The photosynthetic pig-
ments of leaves were estimated according to METZNERet aJ. (1965).
The second group of seedlings was transplanted in the soil of the botanical garden,
Botany Department, Faculty of Science, Tanta University. The plants were cultivated in
randomised blocks, replicated twice and left to grow until the fruiting stage. Air-dried
roots, shoots and fruits were separated, stored in paper bags then dried again in an oven at
40 °C to constant weight (7 days). Thereafter, weights (g) and the numbers of fruits and
seedswere recorded.
The concentrations of sodium, potassium, calcium, magnesium and phosphorus were
determined in roots and shoots according to ALLEN et aJ.(1974). Soluble and insoluble ni-
trogen contents were determined by the method of NAGUm(1969), carbohydrates were es-
timated according to NELSON(1944) and totally soluble proteins were estimated quantita-
tively using LOWRY'SMethod (LoWRY 1951).

HPLC analysis for the plant honnone componentsin algal extracts

Isolation, identification and quantification of plant hormones in the algal extracts were car-
ried out in laboratories of JHG Analytical Service Ltd. Waterford, Ireland on the basis of
Arnerican Society of Testing and Materials D5836 (ASTMD). Analytical Standard Refer-
ence (ASTM!ISO method D 5836) for HPLC analysis was applied by means of A HP-
1050 HPLC (Hewlett Packard) with a gradient solvent delivery system connected to a su-
pelcosil LC-18-DB (25x4.60 mm ID, 0.5 ~m) colurnn. A linear gradientofsolventA (wa-
ter!acetonitrile (90 : 10) in solvent B (acetonitrile) was prograrnmed as the mobile phase
with flow rate of 1.5 ml. min-I Solvent B was maintained at 20 % for 5 minutes after sam-
ple injection and increased gradually until 70 % over 30 minutes. The UV detector was set
at 220 nrn.

Statistical Analysis
To investigatethe relationships betweenconcentrationsof algal extracts and different
growth pararneters,contentsof chernicalcomponentsandyield of Viciafaba at different
growth stages,the Pearsoncorrelationcoefficient (r) was calculatedat 0.05, 0.1 and 0.5
probability levelsandthe testof significancewasperformed.
The one way analysis of variance(ANOVA) was performedto assessthe degreeof
variation betweenconcentrationsof algal extractsand plant variables.The analysiswas
performedat the aboveP levelsof significance.
216 A. H. EL-NAGGAR et a1.

TableI: Effect of seedsoakingin different concentrationsof aqueousextractsof Ulva lactuca on


germinationpercentage,radicaland plumule lengthsand numberof lateral rootsof Viciataba
seedlings.
Gerrnina- Growth Control
tion period Pararneters 0.4% 0.6% 0.8% 1% 1.2% 1.4% F

2 days Genniation [%] 90 100 100 100 100 100 100 0.714...
Radicallength [cm] 0.1 4.03 4.07 4.6 4.7 4.09 3.17 0.373
Plumulelength [cm] - 0.85 1.1 0.7 0.65 0.114
No. or lateral roots - - -

Gennination[%] 100 100 100 100 100 100 100 -


Radicallength [cm] 0.63 5.07 5.44 5.59 5.77 5.01 4.58 0.426
Plumulelength [cm] - 1.59 1.76 2.2 2.9 2.41 1.44 1.648".
No. of lateralroots - 5 6 9 10 6 2 0.116

Germination[%] 100 100 100 100 100 100 100 -


Radicallength [cm] 5.13 6.06 8.2 9.07 10.5 5.36 3.56 0.098
Plumulelength [cm] 3.3 5.15 5.77 7.07 8.2 5 1.7 0.038
No. of lateral roots 6 8 9 9 10 7 3 0.497

Results and discussion


Germination stage

The present investigation showed that the two alga1 extracts increased the per-
centage of germination of '\I: faba seeds. A high1y positive corre1ation was
recorded for germination percentage (after 2 days) in seedstreated with C. kess-
leri extract (Tab1e1 and 2). These resu1tsagree in princip1e with those of MoHAN
et al. (1994), GENCER& AY (1997) and MEHfA et al. (1999) who observed that al-
gal extracts increased germination and seedling growth in several p1ants.
The two algal extracts increased all seed1inggrowth parameters compared to
the control. The most pronounced stimulation was detected in seedlings treated
with 1 % U. lactuca aqueous extract which showed an increase in the radic1e and
p1umule lengths until the 4th dar compared to C. kessleri. However, C. kessleri
extract was more stimulatory for radical and plumu1e 1engthsafter 8 days show-
ing 2.5 and 4.2 fold increase, respective1yin seedlings treated with 1 % (Tab1es1
and 2). Radic1e length of 4-days o1d '\I:faba seed1ingsemerged from seedspre-
soaked in C. kessleri extracts showed highly significant correlation and a high1y
significant F va1ue at P<O.O5. In accordance with the present resu1ts, seaweed
concentrate prepared from Ecklonia maxima (kelpak) was found to increase the
root 1ength and root number of Pinus pinea seed1ings(ATZMONet a1. 1994), in-
creased root and shoot growth in three species of Eucalyptus (VAN STADENet
al.1995) and promoted root formation in a variety of plants (CROUCH& VAN
STADEN1991), which has been attributed to the re1ative1yhigh concentrations of
indo1espresent in the extract (CROUCHet al. 1992). In addition, the stimu1ation of
Inf1uence of Ulva lactuca and Chlorella kessleri on Vicia faba 217

Table2: Effect o• seedsoakingin different concentrationso• aqueousextractso• Chlorella kess-


leri on genninationpercentage,radical andplumule lengthsandnumber o• lateral roots o• Vicia
faba seedlings.
Gerrnina- Growth
tion period Pararneters Control 0.4% 0.6% 0.8% 1% .2% .4% F
2 days Gerrnination[%] 100 100 100 100 100 100 100 0.714***
Radicallength[cm} 0.1 0.15 0.26 0.31 0.68 0.3 0.2 1.946***
Plumule length {cm}
No of lateral roots

4 days Gennination [%] 100 100 100 100 100 100 100
Radica11ength[(cm] 0.63 0.76 1.13 1.15 3.06 2.1 1.98 20.067'
Plumulelength [cm] -
No. of latera1roots -

radicle and plumule lengths and number of lateral roots of ~ faba were reported
by EL-MELEIGY (1999) and EL-SHEEKH & EL-SAlED (2000) using seaweed con-
centrates.
The differential effects of the two algal extracts on the radicle and plumule
lengths may be due to the difference in their hormonal contents especially of
auxin (Fig. 6) which is responsible for enhanced root formation, seed germina-
tion and root and stem elongation (TAKAHASHI 1986), besides cytokinins which
play a role in root deve10pment as suggested by STIRK &VAN STADEN(2001).
Treatment of ~ faba seeds with U. lactuca increased the fresh and dry weights
of roots and shoots up to 0.8 %, whereas higher concentrations were inhibitory.
On the other hand, C. kessleri increased both fresh and dry weights at all concen-
trations. The highest increase in dry weights was by 2.9 and 2.3 f01d of roots and
shoots, respectively, in seedlings presoaked in 1 % C. kessleri extract (Fig. 1).
Several workers have indicated the increase in fresh and dry weights of many
p1ants in respoQse to algal extracts (FEATONBY-SMITH & VAN STADEN 1983,
TAMILSELVAN & KANNAN 1994, VAN STADENet al. 1995, EL-MELElGY 1999). The
increase of fresh and dry weights of ~ faba by the two algal extracts can be re-
ferred to the increase of nutrients uptake and growth enhancement due to the
presence of hormones in both algal extracts which has been reflected to increase
biomass (MOONEY & VAN STADEN 1986, GLASS 1989).
Soaking of ~ faba seeds in U. lactuca and C. kessleri aqueous extracts in-
creased the leaf area of seedlings over the control up to 1 %. However, higher
concentrations (1.4 %) of both extracts reduced leaf area compared to control
(Fig.2). The increase of leaf area might be referred to improving growth and
218 A. H. EL-NAGGAR et a1.

A B

o 02 04 06 08 1 12 14
Co,oo,""", I"') 01 Ul""""" .."~".od'-

Concentration ('!\o) cf U/va lactuca


aqueous extract

Fig. 1. Effect of seedsoakingin different concentrationsof aqueousextractsof Ulva


lactuca (A) and Chlorella kessleri (D) on fresh weight and dry weight) of Viciafaba
seedlings.

vigour of the plant and increasing the uptake of nutrients essential for grOWthdue
to the presence of hormone s as suggested by HaNG et al. (1995), STIRK& VAN
STADEN(1997a, b), especially cytokinin (TAKAHASHI1986) which brought about
a promotion of leaf growth and apical dominance. Similar increase in leaf area in
response to seaweedtreatment was detected in Beta vulgaris by FEATONBY-SMITH
& VAN STADEN(1983), in Eucalyptus by VAN STADENet al. (1995), in Phaseolus
lunatus by REnz & TRUMBLE(1996) and in Viciafaba by EL-MELEIGY(1999).
Aqueous extracts of U. lactuca and C. kessleri elevated significantly the pig-
ment content in leaves of ~ faba seedlings. Thus, 1 % of C. kessleri extract in-
duced a remarkable increase in Chl a, Chl b and carotenoids (5.8, 12.8 and 14.2
fold, respectively) compared to control (Fig. 2). Similar results were obtained by
WHAPHAMet al. (1993), EL-MELEIGY(1999) and EL-SHEEKH& EL-SAIED(2000).
The increase in chlorophyll in response to algal extracts treatrnent can be referred
to the role of algal concentrate in enriching magnesium, thus stimulating its avail-
Influenceof Ulva lactucaand Chlorellakesslerion Viciafaba 219

A B

15

~E 10
~
~
~ ,,
;
~
~
5 ~
..J

o
o 0.2 0.4 0.6 0.8 1 1.2 1.4
Concerdration (%)of
Chlonl'. ke..le,-;-queous

o 0.2 04 0.6 08 1 12 14

ConcOnlr3ion (~) oj OIobfPNa kossion


aquoous oxtr3ct

Fig. 2. Effect of seedsoakingin different concentrationsof aqueousextractsof Ulva


lactuca (A) and Chlorella kessleri(B) on leaf areaandphotosyntheticpigmentsof Vicia
faba seedlings.

ability and uptake from both algal extracts and soil as indicated by MOONEY&
VAN STADEN(1986). Moreover, the enhancementof chlorophyll formation CaDbe
due to the effect of cytokinin (TAKAHASHI1986) and betaines (WHAPHAMet al.
1993, BLUNDENet al. 1997) present in algal extracts as indicated in the present
study ( Fig.6).

Fruiting stage
Inorganic elementsin roots and shoots

The inorganic elements, including Na+, K+, Mg2+, Ca2+ and ps+, in roots and
shoots of harvested ~ faba presoaked in U. lactuca and C. kessleri aqueous ex-
tracts were usually higher than the controls (Table 3). However, C. kessleri was
more stimulatory for the accumulation of Na+ and K + in roots and shoots. A con-
220 A. H. EL-NAGGAR et al.

Table3: Effect of seedsoakingin different concentrationsof aqueousextractsof Ulva lactuca and


Chlorellakesslerion contentsof inorganicelementsof harvestedViciafaba plants(fruiting stage).

~
ti Conc. Na+[mg.g-l] K+ [mg.g-l] Mg2+[mg.g-l] Ca2+[mg.L-l] p5+ [mg.g-I]
K
" [%]of
~ the
O
"C' extract Root Shoot Root Shoot Root Shoot Root Shoot Root Shoot
«

centration of 1.2 % of C. kessleri resulted in 7.8 and 9.3 fold increase in K+ in


roots and shoots of ~ faba, respectively, compared to the control. On the con-
trafY, U. lactuca extract increased magnesium, calcium and phosphorus com-
pared to C. kessleri and the stimulatory effect of U. lactuca was more obvious in
roots than in shoots. The positive effect of algal extracts on the inorganic ele-
ments in plants caD be due to the promotion of nutrient uptake from soil
(MOONEY& VAN STADEN1986) supplying the plants with minera! elements such
as phosphorus and ca!cium (DEVILLIERSet aJ. 1983), potassium (BECKETT& VAN
STADEN1989), magnesium (EL-SHEEKH& EL-SAIED 2000) and zinc (BECKETT
&VAN STADEN1990 b). Increasing evidence exists from the raci that nutrient up-
take and movement within plants is under hormona! control (GLASS1989).

Total organic nitrogen in roots and shoots

In genera!, the nitrogen contents (soluble and insoluble) of plants emerged from
seeds soaked in the two algal extracts were high compared to control plants.
However, U. lactuca extract was more stimulatory for the accumulation of solu-
ble nitrogen in roots and insoluble nitrogen in roots and shoots than C. kessleri
extract. The latter extract produced higher soluble nitrogen contents in shoots
than the former ODe.The two alga! extracts produced higher accumulation of in-
soluble nitrogen in shoots than in roots (Fig. 3). This can be attributed to the fact
Inf1uence o• Ulva lactuca and Chlorella kessleri on Vicia faba 221

Fig. 3. Effect of seedsoaking in different concentrationsof aqueousextractsof Ulva


lactuca (A) and Chlorella kessleri(B) on nitrogencontentsof Viciafaba plants(fruiting
sta~e).

lhal the algal extracts can stimulate the translocation of nitro gen from roots to
shoots.
The stimulating effect of algal extracts on nitrogen accumulation can be due to
their promotion of nitrogen uptake from the soil as concluded by BECKE1T& VAN
STADEN(1990 a). Furthermore, algal concentrates which are rich in inorganic
compounds could supply plants with nitrogen. In this regard, TEMPLEet al.
(1989) found that application of kelps (Macrocystis integrifolia and Ecklonia
maxima) as foliar spray increased the growth andnitrogen content of Phaseolus
vulgaris.

Protein and carbohydrates of Viciafaba pods

Application o• 1 % o• each U. lactuca and C. kessleri extract increased pod pro-


teins o• Viciafaba by 57 and 44%, respectively, (Fig. 4).
C. kessleri extract was more effective in accumulation o• direct reducing sug-
ars compared to U. lactuca extract. The latter showed an increase in tota1 reduc-
ing sugars more than C. kessleri extract. A level o• 1 % C. kessleri extract
increased both direct and total reducing va1ues2.2 •old. These results are in ac-
222 A. H. EL-NAGGAR et aj.

A B

45 - ---:
40 J
~ 35.
1RV F~!
01
g 30
.,
" 25
~
"tJ 20
.§ 15
.o
~ 10.
U

o
o 02 0.4 0.6 0.8 1 1.2 1 4
Concentrmion (%) 01 Cfllorella ke.s.sleri
aqueous exlracl

Fig. 4. Effect of seedsoaking in different concentrationsof aqueousextractsof Ulva


lactuca (A) and Chlorella kessleri (B) on protein and carbohydratesof fruits of Vicia
faba plants(fruiting stage).

cordance with BLUNDENet al. (1979) who concluded that foliar application of
aqueous seaweed extracts produced a significant increase in the carbohydrate
content of sugar beets. In addition, EL-SHEEKH& EL-SAIED(2000) reported an in-
crease of proteins and carbohydrates in ~ faba treated with crude extracts of
some green and red seaweeds.
The increase in the protein and carbohydrate contents in response to U. lac-
fuGaand C. kessleri aqueous extracts could be due to the presence of cytokinin in
these algal extracts which promote protein synthesis and cen division (POZSARet
al. 1967). Hence, seaweed extracts may play a direct role in protein synthesis as
suggested by NoRRIE& H!Ln (1999). Moreover, accumu1ation of carbohydrate
cou1d induce an increase in respiration rate accompanied with a rise in a-keto
acids concentrations. These acids are required for the biosynthesis of different
amino acids needed for protein biosynthesis.
Influenceof Ulva lactuca and Chlorella kesslerion Vicia.faba 223

Yield
AII concentrations of the two algal extracts increased the yield parameters of
~ faba. The highest oDe was detected with 1 % of U. lactuca extract which in-
creasedthe weight and number of fruits, weights and number of seedsin all fruits
of ~ faba over the control by 4.4, 3.9, 5 and 3.9 fold, respectively (Fig. 5). How-
ever, 0.6 % of C. kessleri extract induced a 3, 3.2, 1 and 3.2 fold increase in the
same parameters, respectively. As a general trend, plants treated with U. lactuca
extract gave higher yields than those treated with C. kessleri extract.
Several authors reported on the beneficial effects of algal extracts as natural
regulators in increasing the yields of many crops. In this concem, application of
seaweed extracts increased significantly the yield of potatoes (BLUNDENet al.
1979, FEATONBY-SMITH & VAN STADEN1983 and LOPEZ-MosQUERA& PAZOS
1997), wheat (BECKETT& VAN STADEN1989), bean (TEMPLE& BOMKE1989) and
Ziziphus mauritiana (AMA RAO 1991). Furthermore, NORRlE& HILTZ (1999)
recorded positive results from the application of seaweed extract (Ascophyllum
nodosum) on the yield of many crops. The positive influence of algal extracts on
the yield of treated plants can be attributed to their contents of hormones, espe-
cially cytokinin, auxin and gibberellin (TEMPLE& BOMKE1989).

The content of hormones in algal extracts

Four plant hormones have been identified in the extracts of U. lactuca and
C. kessleri, namely: cytokinin, auxin, gibberillin and betaines (Fig. 6). However,
the content of cytokinin and betaines in U. lactuca (2.35 and 25.1 ~g .kg-l dry
weights, respectively) were higher than those in C. kessleri (1.89 and 21.6
~g. kg-l dry weights, respectively). On the other band, C. kessleri extract po s-
sessed higher levels of auxin and gibberillin (8.15 and 11.90 ~g.kg-l dry
weights, respectively) than U. lactuca which contained 6.65 and 10 ~g. kg-l dry
weights, respectively.
Most crop responses to a1ga1extracts application have been considered as be-
ing primarily due to the cytokinin and auxin compounds (CROUCH 1991, STlRK &
VAN STADEN 1997a, b, EL SHEEKH & EL SAIED 2000). The magnitude of the
growth responses following seaweed application suggests an additive effect of
both enhanced nutrient uptake (CROUCH et al. 1990) and regulatory action of
plant growth substances.
Identification of cytokinins (STEPHEN et a1. 1985, MOONEY & VAN STADEN
1986, STIRK &VAN STADEN 1997 a, b, EL-MELEIGY 1999 & EL SHEEKH & EL
SAlED 2000), auxin (TEMPLE & BOMKE 1989, CROUCHet a1. 1992, CROUCH& VAN
STADEN 1993, HONG et a1. 1995), gibberellin (GOPALA 1984 and CROUCH 1991)
and betaines (WHAPHAMet a1.1993, TYrnAK et a1.1994,Wu et a1. 1997) in severa1
commercia1 seaweed concentrates implied their possible involvement in the stim-
ulatory effect such as found in the present investigation. In addition, cytokinin
and auxin were identified in seven strains of unicellular Chlorophyta and three
strains of Cyanophyta (STlRK et a1. 2002).
224 A. H. EL-NAGGAR et al.

o 0.2 0.4 0.6 0.8 1 1.~ 1.4

140-.
120 F=O.321 ,

:9 100.
1:-
~
.. 80-
3
'" 60-
~
:;., 40.
20
O
o 0.2 0.4 0.6 0.8 1 1.~ 1.4

250
200 F=O.5007 --,

150
'"
"Q 100
..
..
(/)
50

O
o 0.2 0.4 0.6 0.8 1 1.2 1.4

Concentration (~) 01 U/va /act/ICa


aqueous eX!~ct

Fig. 5. Effect of seedsoaking in different concentrationsof aqueousextractsof Ulva


lactuca (A) and Chlorella kessleri (B) on yield of harvestedViciafaba plants (fruiting
stage).
Influence of Ulva lactuca and Chlorella kessleri on Vicia faba 225

30

25

20
CI)
Q)
c
o
E
...
o
.c
-O 15
...
C
OJ
...
C
O
U

10

o
2 3 4
Hormones

Fig. 6. Plant growth hormonesof Ulva lactucaand Chlorella kessleri.

The present study favours again the use of algal aqueous extracts as a safe
agricultural fertilizer for improving the quality and yield of mailY crops on a large
scale.
226 A. H. EL-NAGGAR et a1.

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228 A. H. EL-NAGGAR et aJ.

Manuscript received May 27, 2004, accepted March 3, 2005.

~
Influenceof Ulva lactucaand Chlorella kesslerion Viciafaba 229

J Authors' address:

AMAL H. EL-NAGGAR,
MoHAMED E.H. OSMAN,
MOSTAFA M. EL-SHEEKH,
j SALV F. GHEDA
TantaUniversity
Faculty of Science,
BotanyDepartment
Tanta,Egypt.

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