J Appl Phycol (2014) 26:619–628

DOI 10.1007/s10811-013-0078-4

Effect of liquid seaweed extracts on growth of tomato

seedlings (Solanum lycopersicum L.)
Rosalba Mireya Hernández-Herrera &
Fernando Santacruz-Ruvalcaba & Mario Alberto Ruiz-López &
Jeffrey Norrie & Gustavo Hernández-Carmona

Received: 12 March 2013 / Revised: 26 June 2013 / Accepted: 27 June 2013 / Published online: 17 July 2013
# Springer Science+Business Media Dordrecht 2013

Abstract Seaweed extracts are used as nutrient supplements, receiving LSEs of U. lactuca and P. gymnospora showed
biostimulants, or biofertilizers in agriculture and horticulture increased shoot length, root length, and weight. Furthermore,
to increase plant growth and yield. In this study, we examined U. lactuca and P. gymnospora were found to be more suc-
the effect of liquid seaweed extracts (LSEs) made from Ulva cessful and better candidates for developing effective
lactuca, Caulerpa sertularioides, Padina gymnospora, and biostimulants to improve the growth of tomato plants. This
Sargassum liebmannii as biostimulants on the germination study provides important information on the identification and
and growth of tomato (Solanum lycopersicum) under labora- utilization of Mexican seaweed resources for agriculture and
tory and greenhouse conditions using foliar and soil drench is the first study to report on the uses of these seaweeds as a
applications of LSEs. We assessed LSEs at different concen- source of liquid extracts as biostimulants in agriculture.
trations (0.2, 0.4, and 1.0 %) on germination parameters
(percentage, index, mean time, energy, and seedling vigor Keywords Seed germination . Seaweeds . Extract . Nutrient
index) and growth parameters (plumule length, radical length, analysis . Biofertilizer . Chlorophyta . Phaeophyta
shoot length, root length, fresh weight, and dry weight) of
tomato seedlings. Our results indicate that seeds treated with
LSEs of U. lactuca and P. gymnospora at lower concentra- Introduction
tions (0.2 %) showed enhanced germination (better response
in germination rate associated with lower mean germination Seaweed and seaweed products have been used worldwide to
time, high germination index and germination energy, and increase plant growth and yield. Modern agriculture is
consequently greater seedling vigor and greater plumule and searching for new biotechnologies that would allow for a
radicle length). Application as a soil drench was found to be reduction in the use of chemical inputs without negatively
more effective in influencing the height of the plant (up to affecting crop yield or the farmers' income. In recent years,
79 cm) than the foliar spray application (75 cm). Plants the use of natural seaweed as fertilizer has allowed for partial
substitution of conventional synthetic fertilizer (Dhargalkar and
Pereira 2005; Hong et al. 2007; Khan et al. 2009; Zodape et al.
R. M. Hernández-Herrera : F. Santacruz-Ruvalcaba :
2010). In addition, a number of commercial seaweed extract
M. A. Ruiz-López
Centro Universitario de Ciencias Biológicas y Agropecuarias, products are available for use in agriculture and horticulture
Universidad de Guadalajara, Carretera Guadalajara-Nogales Km and can be used as liquid extracts applied as foliar spray, soil
15.5, C.P. 45110 Zapopan, Jalisco, Mexico drench, or in granular/powder form as soil conditioners and
manure (Blunden et al. 1997; Lingakumar et al. 2004;
J. Norrie
Acadian Seaplants Limited, Dartmouth NS B3B 1X8, Canada Thirumaran et al. 2009). These extracts are marketed as liquid
biostimulants because a chemical analysis of seaweeds and
G. Hernández-Carmona (*) their extracts has revealed the presence of a wide variety of
Instituto Politécnico Nacional–Centro Interdisciplinario de
plant growth-promoting substances such as auxins, cytokinins,
Ciencias Marinas, Av. Instituto Politécnico Nacional s/n, Col. Playa
Palo de Santa Rita, C.P.23096 La Paz, Baja California Sur, Mexico and betaines (Khan et al. 2009). These substances can influence
e-mail: [email protected] shoot and root system development (Durand et al. 2003; Stirk
620 J Appl Phycol (2014) 26:619–628

et al. 2004). As well, macronutrients and micronutrients can absorption spectrophotometry for mineral analysis of sodi-
help promote the growth of various vegetables, fruits, and other um, potassium, calcium, and phosphorus (by colorimetry)
crops (Blunden 1991; Crouch and van Staden 1993; Moller and following procedures from the Association of Official Ana-
Smith 1998). Many beneficial effects have been reported on the lytical Chemists (AOAC 1990). Then, 100 g of each sample
use of seaweed extracts. Positive responses include improved was added to 1 L of distilled water with constant stirring for
germination, root development, leaf quality, general plant vig- 15 min followed by autoclaving at 121 °C for 1 h at 1.21 kg-
or, and resistance to pathogens (Khan et al. 2009). cm−2. The hot extracts were filtered through a Whatman No.
Seaweed is an inexpensive local resource along coastal 40 filter paper and stored. The liquid seaweed extracts
agricultural areas. In Mexico, seaweeds are abundantly avail- (LSEs) were designated as stock solution and coded
able and represent a great potential for eventual commercial according to the genus and species: U. lactuca (UL), C.
exploitation (Gojón-Báez et al. 1998). However, the only sertularioides (CS), P. gymnospora (PG), and S. liebmannii
commercial extracts currently produced in Mexico from raw (SL). Furthermore, the pH and electrical conductivity (EC)
seaweeds are Kelpro and Kelprosoil from Macrocystis pyrifera of the LSEs were measured using a pH meter and conduc-
(SAGARPA 2012) and Algaenzims from Sargassum spp. tivity meter, respectively, and the latter is expressed as dS
(Canales-López 2000; Sunarpi et al. 2010). These products m−1. Finally, the color of the seaweed extracts was ob-
are available for use in agriculture and horticulture. One pos- served visually. All determinations were performed in
itive step towards the inclusion of native seaweed resources in triplicate.
Mexico is to use biofertilizers derived from seaweed as an
alternative input to improve negative cropping conditions such Bioassay for germination test
as the progressive degradation of ecosystems and the contam-
ination of agricultural lands caused by synthetic fertilizers. Experiments were conducted using certified tomato seeds (S.
Tomatoes (Solanum lycopersicum) are one of the most lycopersicum cv. Rio Fuego; Cal-Oro Vegetable Seeds, United
important vegetable crops around the world in terms of human Genetics, Inc., USA). Germination was observed daily over a
consumption, and they are also the most popular garden veg- period of 8 days according to methods of the Association of
etable. In 2009, Mexico was among the top ten tomato- Official Seed Analysts (AOSA 2005). Four groups of 100
producing countries contributing with almost three million seeds were tested for germination per treatment (AOSA
tonnes (FAO 2009) to the global production. However, one of 2005). Experimental units were arranged in a randomized
the main problems facing tomato production in Mexico is the complete block design. Before treatment with LSEs, tomato
intensive application of chemical fertilizers causing damage to seeds were surface-sterilized in 4 % sodium hypochlorite
the soil ecology and agricultural systems (Villarreal-Sánchez solution for 10 min and subsequently triple-rinsed in sterile
et al. 2003). The aim of this study was to examine the effect of distilled water prior to soaking in the seaweed extracts. Tested
liquid seaweed extracts derived from Mexican resources on tomato seeds were placed on a Whatman No. 5 filter paper in
seed germination and growth of tomato plants. sterilized 90-mm Petri dishes and then treated with 5 mL
distilled water (control) or different concentrations of LSEs
(0.2, 0.4, and 1.0 %). The plates were incubated at 25±1 °C
Materials and methods and 16-h light/8-h dark regime. Germination was considered to
have occurred once the radicle had protruded more than 2 mm.
Preparation of liquid seaweed extracts Measured variables included germination percentage (GP),
germination index (GI), mean germination time (MGT), and
Four algal species, two green, Ulva lactuca Linnaeus and germination energy (GE), as well as seedling vigor index
Caulerpa sertularioides Gmelin, and two brown, Padina (SVI), plumule length, radicle length, total plant height, and
gymnospora (Kützing) Sonder and Sargassum liebmannii dry weight of tomato seedlings. After 12 days, the LSEs effects
J. Agardh seaweeds, were collected from the intertidal zone on seed germination and growth of tomato seedlings were
at low tide, in May and November 2009, from the coastal measured. Plumule length, radicle length, and total plant height
area of Jalisco, Mexico, in Bahia Tenacatita (19° 28″N, 104° were measured with a vernier caliper. Dry weight was obtained
84″W) and Bahia Careyitos (19° 43″N, 105° 02″W). Sea- with an electronic balance after oven-drying at 60 °C.
weed species were collected by hand and washed with sea- Parameters were calculated as follows: GP=(number of-
water to remove debris, shells, and sand. Samples were germinated seeds/total number of seeds)×100. GI was calcu-
transported to the laboratory in plastic bags, washed with lated as described by the Association of Official Seed Analysts
tap water to remove surface salt, oven-dried for 72 h at 60 °C, (AOSA 1983), following the equation: GI=∑(Gt/Tt), where
and then ground in an electric mill (IKA-M 20) to less than Gt is the number of seeds germinated on day t and Tt is the
0.50 mm. This milled material (100 g) of each sample was number of days. MGT was estimated according to Ellis and
subjected to acid digestion and analyzed by atomic Roberts (1981) and expressed as days. MGT=∑(D×n)/∑n,
J Appl Phycol (2014) 26:619–628 621

where n is the number of seeds germinated on day D and D is ANOVA was carried out for each parameter studied to assess
the number of days counted from the beginning of the significant differences. In the second experiment, three-way
test. Seed GE was calculated according to the formula ANOVA assessed significant differences at the 5 % level. All
(GE %)=(number of germinating seeds/number of total statistical analyses were performed with Statistical Package
seeds per treatment after germination for 3 days)×100. STATGRAPHICS® Centurion XV for Windows.
SVI was determined according to Orchard (1977) by the
following formula: SVI=(seedling length (cm)×germination
percentage). Results

Greenhouse growth bioassay Macroelements in seaweeds and physicochemical content

of LSEs
Tomato plants were grown in a growth chamber under 16-h
light regime at 25 °C and 8-h dark regime at 18 °C in sterilized Results obtained from the nutrient analysis showed the presence
soil peat moss (Sunshine Mix 3™). Two hundred fifty 15-day- of the macroelements Na, K, Ca, and P in all samples. The
old plants were selected and randomly assigned to different concentration of Na was higher in the green seaweeds U. lactuca
treatment groups and transplanted into pots containing a sand, and C. sertularioides, but K concentration was higher in the
perlite (Termolita S.A., Mexico), volcanic rock, and peat moss brown seaweeds P. gymnospora and S. liebmannii. The concen-
(Sunshine Mix 3™) (1:1:1:1 w/w) soil mix. Plants were fer- tration of Ca was higher in P. gymnospora and C. sertularioides.
tilized 1 week after transplanting and treated with 50 mL of The P concentration was low in all seaweeds (Table 1). The pH
20:20:20 (N–P–K) soil drench solution (Peters Professional; values for LSEs of U. lactuca and P. gymnospora were neutral
Scotts-Sierra Horticultural Products Co., USA). Thereafter, and slightly acidic than those for LSEs made from C.
the plants were irrigated using the LSEs (50 mL every week). sertularioides and S. liebmannii. The value of EC increases in
Plants were also irrigated separately with water (50 mL every all LSEs with an increase in the concentrations (Table 2).
third day). Potted plants were grown for 7 weeks in a green-
house at ~25±2 °C, in 85 % relative humidity. Morphological Effect of LSEs on seed germination and growth of tomato
characteristics such as shoot length, root length, total height, seedlings under laboratory conditions
fresh weight, and dry weight were measured.
The experimental design and treatment were similar to those Germination occurred in all treatments after 2 days. The effect
reported by Crouch and van Staden (1992). A total of 25 of the liquid seaweed extracts of U. lactuca and P. gymnospora
different treatments were used with ten replications. The first at a concentration of 0.2 % gave a significant (P≤0.05) in-
treatment served as the control in which plants were grown crease in GP over control after 2 days (Fig. 1a). Data indicated
without LSEs. Three factors were randomized for the other 24 that the same treatments at concentrations of 0.4 and 1.0 %
LSE treatments. The first factor was LSE applications method decreased the GP (Fig. 1b, c). Treatments of C. sertularioides
(foliar spray versus soil drench). The second factor was the and S. liebmannii had an inhibitory effect on seed germination.
type of seaweed species used to produce the LSEs. The third All concentrations delayed germination and GP dropped off
factor was the concentration (0.2, 0.4, and 1.0 %). The exper- with a higher concentration (1.0 %) (Fig. 1c).
imental units were arranged in a completely randomized tri- Seeds treated with U. lactuca and P. gymnospora LSEs
factorial design receiving either 50 mL distilled water for the showed higher germination rate associated with lower MGT
control or 50 mL LSEs for the experimental treatments. and greater seedling vigor (Table 3). LSEs of U. lactuca and P.
gymnospora at 0.2 % showed high GP (75 and 76 %, respec-
Statistical analysis tively), elevated GI (9.8 and 10, respectively), a reduction in
MGT (5.6 days), an increase in GE (88.7 and 87.2 %, respec-
All data were analyzed for significant differences by analysis tively), and enhanced SVI (1,026.7 and 1,262.3, respectively).
of variance (ANOVA) with mean separation using least sig- In contrast, seeds treated with LSEs of C. sertularioides and S.
nificant difference (LSD). In the first experiment, one-way liebmannii at 1.0 % had the longest average delay (high

Table 1 The content of

macroelements in seaweed spe- Species Na K Ca P
cies (g·100 g−1, dry wt.)
U. lactuca 5.57±0.80 1.85±0.30 1.88±0.06 0.10±0.08
C. sertularioides 4.42±0.40 0.47±0.40 3.10±0.50 0.20±0.08
P. gymnospora 1.81±0.50 4.27±0.60 3.65±0.40 0.10±0.08
Values are average ± standard S. liebmannii 1.56±0.40 4.54±1.00 1.85±0.08 0.17±0.05
error (n=3)
622 J Appl Phycol (2014) 26:619–628

Table 2 Physicochemical con-

tent of liquid seaweed extracts Treatments (%) Color pH EC (dS m−1)
(LSEs) treatments of U. lactuca
(UL), C. sertularioides (CS), P. Control 7.0 0
gymnospora (PG), and S. UL 0.2 7.00±0.50 0.99±0.11
liebmannii (SL) UL 0.4 Greenish yellow 7.30±0.50 1.74±0.11
UL 1.0 7.36±0.50 3.57±0.11
CS 0.2 6.71±0.50 1.01±0.20
CS 0.4 Dark green 6.96±0.50 1.85±0.20
CS 1.0 6.99±0.50 3.93±0.20
PG 0.2 7.10±0.20 0.77±0.10
PG 0.4 Brownish red 7.60±0.20 1.43±0.10
PG 1.0 7.60±0.20 2.98±0.10
SL 0.2 6.19±0.50 1.01±0.15
Values are average ± standard SL 0.4 Brown 6.26±0.50 2.61±0.15
error (n=3)
SL 1.0 6.37±0.50 3.99±0.15
EC electrical conductivity

100 a MGT), were the latest to germinate, and had the greatest
90 spread of germination over time (Table 3).
80 Control
Germination (%)

70 UL The LSEs had a significant effect (P≤0.05) on growth of

60 CS tomato seedlings. All LSEs showed a stimulatory effect on
50 PG plumule length. The highest average plumule length was
40 SL
30
found in plants that received LSEs from U. lactuca and P.
20 gymnospora at 1.0 % (7.3 and 8.3 cm, respectively; Fig. 2a).
10 Moreover, these results indicate that LSEs of U. lactuca, P.
0
1 2 3 4 5 6 7 8 gymnospora, and S. liebmannii promoted radicle length. The
Days after sowing highest average radicle length was found in plants that received
100 b LSEs from U. lactuca, P. gymnospora, and S. liebmannii at
90 0.2 % (4.1, 5.8, and 5.5 cm, respectively) in comparison to the
80
Germination (%)

70
control. Furthermore, all C. sertularioides treatments had an
60 inhibitory effect on radicle length (2.2 to 2.8 cm, Fig. 2b). All
50 C. sertularioides and S. liebmannii LSE treatments significant-
40
30
ly increase dry weight of tomato plants, as did U. lactuca and P.
20 gymnospora LSEs at (1.0 %). The highest dry weight (0.036 g)
10 was higher for the brown algae LSEs (Fig. 2c).
0
1 2 3 4 5 6 7 8
Days after sowing Effect of LSEs on tomato seedling growth in greenhouse

100 c In the greenhouse experiment, both foliar spray and soil

90
80 drench of LSEs showed a significant enhancement (P≤0.05)
Germination (%)

70 on growth of tomato plants. Application as a soil drench was

60 found to be more effective in influencing the height of the
50
40 plant (up to 79 cm) than the foliar spray application (75 cm).
30 The interaction between application form, treatment, and
20 concentration demonstrates that plants treated with foliar sprays
10
0 of U. lactuca at 1.0 % and P. gymnospora at 0.2 % displayed an
1 2 3 4 5 6 7 8 increase in shoot length (47 and 49 cm, respectively), but plants
Days after sowing receiving the same extracts as soil drench showed no significant
Fig. 1 Germination percentage of tomato seeds treated with liquid difference in shoot length (Fig. 3a, b). Treatments applied as
seaweed extracts at concentrations of a 0.2 %, b 0.4 %, and c 1.0 %
foliar spray of U. lactuca and P. gymnospora at 0.2 % displayed
from U. lactuca (UL), C. sertularioides (CS), P. gymnospora (PG), and
S. liebmannii (SL). Values are presented as average (n=400 seeds); bars an increase in root length (25 and 26 cm, respectively) and total
represent standard error plant height (70 and 75 cm, respectively). Similarly, soil drench
J Appl Phycol (2014) 26:619–628 623

Table 3 Effects of liquid seaweed extracts (LSEs) treatments on germination parameters of tomato seeds: germination percentage (GP), germination
index (GI), mean germination time (MGT), germination energy (GE), and seedling vigor index (SVI)

LSEs (%) Parameters

GP GI MGT (days) GE (%) SVI

Control 31±1.40 c 8.9±0.24 f 5.9±0.08 ef 68.8±2.76 d 763.4±40.8 b

UL 0.2 75±2.00 g 9.8±0.89 hi 5.6±0.06 a 88.7±2.76 e 1,026.7±40.8 ef
UL 0.4 55±1.85 e 9.4±0.86 gh 5.8±0.08 b 74.8±2.76 d 946.8±40.8 de
UL 1.0 50±1.62 de 5.2±0.54 b 5.8±0.11 bc 72.9±2.76 d 1,096.5±40.8 fg
CS 0.2 1±0.60 a 8.2±0.72 e 6.3±0.09 h 49.8±2.76 c 805.7±40.8 bc
CS 0.4 12±3.93 b 9.0±0.93 fg 6.2±0.24 g 56.9±2.76 c 891.6±40.8 cd
CS 1.0 5±0.91 a 3.4±0.75 a 6.5±0.10 i 24.5±2.76 a 622.9±40.8 a
PG 0.2 76±1.10 g 10±0.67 i 5.6±0.06 a 87.2±2.76 e 1,262.3±40.8 h
PG 0.4 66±1.67 f 9.9±0.57 hi 5.7±0.05 a 84.5±2.76 e 1,229.7±40.8 h
PG 1.0 46±2.10 d 9.1±0.92 fg 5.8±0.09 bcd 74.2±2.76 d 888.7±40.8 cd
SL 0.2 1±0.40 a 7.9±0.85 de 5.9±0.13 def 31.3±2.76 ab 1,160.6±40.8 gh
SL 0.4 2±0.57 a 7.5±1.02 d 5.9±0.16 cde 36.0±2.76 b 873.2±40.8 bcd
SL 1.0 1±0.31 a 6.6±1.13 c 6.0±0.18 f 28.5±2.76 ab 635±40.8 a

Values are average ± standard error (n=400). Average followed by the same letter within columns is not significantly different, according to LSD
multiple range test (P≤0.05)
UL U. lactuca, CS C. sertularioides, PG P. gymnospora, SL S. liebmannii

applications of U. lactuca at 1.0 % and P. gymnospora at 0.2 % and neutral extracts can affect bioactivity (Booth 1969; Henry
resulted in an increase in root length (33 and 32 cm, respec- 2005). According to Reinhardt and Rost (1995), most plants
tively) and total plant length (79 cm) (Fig. 3c–f). are more sensitive to salinity during germination and seedling
In addition, positive effects on fresh weight were observed growth. This is in agreement with our study whereby a higher
with application as foliar spray of P. gymnospora at all concen- germination percentage was found with treatments of U.
trations (9.9, 10.1, and 10.7 cm) and with application as soil lactuca and P. gymnospora (0.99 and 0.77 dS m−1 with
drench of U. lactuca and C. sertularioides at 1.0 % (14 and 0.2 %) at low concentration. This may be due to the absence
14.6 cm, respectively) (Fig. 4a, b). Dry weight of tomato plants of salts in the medium, thereby allowing seeds to more effi-
was unaffected by treatment with LSEs, except for S. liebmannii ciently imbibe water. In contrast, LSEs of C. sertularioides and
that produced negative effects (Fig. 4c, d). S. liebmannii (3.9 dSm−1 with 1.0 %) at higher concentration
showed a significant negative effect on the germination of
tomato seeds, by inhibiting the seeds' ability to imbibe water.
Discussion Similar results have been reported by Basher et al. (2012).
They evaluated the effect of seaweed, salt water, and drainage
Previous papers reported that seaweeds contain high macro- water on germination percentage, rate, and seedling growth of
element levels (Ca, K, P), especially those from the Phaeophyta tomato (Lycopersicum spp.). The combination of water with
(Hong et al. 2007). The content of minerals in the seaweeds used seaweed treatments shows the highest rate for 2.7 dSm−1 with
in this research was in general agreement with the typical values 0.1 % seaweed concentration and lowest rate for 5 dSm−1 with
for these marine algae from other countries (Gireesh et al. 2011; 0.05 % seaweed concentration compared to control. Salt stress
Hong et al. 2007; Kalaivanan and Venkatesalu 2012; Sivasangari reduced the growth of the plumule and radicle in tomato
et al. 2010) and Mexico (Carrillo-Domínguez et al. 2002; seedlings, and there was a direct adverse relation between NaCl
Robledo and Freile Pelegrin 1997). The quantitative ranges for concentration and reduction in growth (plumule and radicle)
the various components in seaweed can vary due to season, (Nyagah and Musyimi 2009). Tomato seedlings are moderate
location, and analytical methods (Castro-González et al. 1996; sensitive to NaCl salinity. However, during imbibition, the
Ito and Tsuchiya 1981). effect of salt is merely osmotic until a hydration threshold is
The pH and EC from LSEs affected germination in tomato. surpassed (Almodares et al. 2007). Tomato seeds treated with
The beneficial effects of seaweed extracts may arise from low concentrations of U. lactuca and P. gymnospora responded
higher seed moisture after the drying phase (Weges and better in terms of germination rate associated with lower MGT,
Karsssen 1990). Additionally, changes in pH and EC of acidic high GI and GE, and consequently greater seedling vigor and
624 J Appl Phycol (2014) 26:619–628

10 a Super Marmand, and Tanshet Star) at high salt concentration

9 f
de e in proportion to the control application (Al-Harbi et al. 2008).
Plumule length (cm)

8 d de d
c Salt stress decreased germination by preventing the ger-
7 bc b bc b
6 a mination processes due to the accumulation of a high con-
5 a
4
centration of Na+ and Cl− ions, which might be toxic to the
3 embryo or the developing seedlings (Almodares et al. 2007),
2 and can affect many vegetable crops, leading to uneven stand
1
0
establishment and reduction of crop yields (Yildirim and
C UL CS PG SL Guvenc 2006). The mechanisms for salt damage during ger-
Liquid seaweed extracts
mination are not completely understood (Almodares et al.
10 b 2007). However, the effect of salinity on plant growth is a
9
complex syndrome that involves osmotic stress, ion toxicity,
Radicle length (cm)

8
7 h and mineral deficiencies (Musyimi et al. 2007).
6 g
f In the present study, higher concentrations of seaweed ex-
5 e f f
4 e d tract from C. sertularioides, P. gymnospora, and S. liebmannii
3 b c a bc at 1.0 % had toxic effects on tomato seedlings and caused
a
2 detrimental effects such as radicle browning and disintegration
1
0 of plumules. This detrimental effect for tomato growth has been
C UL CS PG SL reported previously by Arnon and Johnson (1942) and is
Liquid seaweed extracts caused by higher pH in the growth medium. Higher concen-
0.05 c trations of seaweed extracts from S. johnstonii (2.0 to 10 %)
caused the same detrimental effects in black gram (Vigna
0.04 cd d d bcd
bcd cd bcd bcd mungo) seedlings (Kumari et al. 2011). Indeed, several studies
Dry weight (g)

ab abc ab ab ab
0.03 a have examined seaweed extracts on seed germination of vari-
ous species such as table beet (Wilczek and Ng 1982), lettuce
0.02 (Moller and Smith 1998), tomato (Demir et al. 2006), green
0.01 gram (Ashok-Kumar et al. 2012), and black gram (Kalaivanan
and Venkatesalu 2012; Ganapathy Selvam et al. 2013). The
0.00 increased germination percentage at low concentrations could
C UL CS PG SL
Liquid seaweed extracts be due to the presence of growth-promoting substances such as
Fig. 2 Effect of liquid seaweed extract treatment applications on a indole acetic acid, indole-3-butyric acid, gibberellins, cytoki-
plumule length, b radicle length, and c dry weight of tomato seedlings nins, micronutrients, vitamins, and amino acids (Challen and
at different concentrations: 0.2 % (white columns), 0.4 % (gray col- Hemingway 1965).
umns), and 1.0 % (black columns). Treatments: control (C), U. lactuca Seaweed products exhibit growth-stimulating activities,
(UL), C. sertularioides (CS), P. gymnospora (PG), and S. liebmannii
(SL). Columns denoted by a different letter are significantly different at and the use of seaweed formulations as biostimulants in crop
P≤0.05. Values represent average (n=300 seedlings); bars represent production is well established. Seaweed ingredients include
standard error macro- and microelement nutrients, amino acids, vitamins,
cytokinins, auxins, and abscisic acid that affect cellular me-
tabolism in treated plants, leading to enhanced growth and
higher plumule and radicle length. The higher concentration crop yield (Crouch and van Staden 1993; Stirk et al. 2004;
showed a decreasing trend, particularly with C. sertularioides Wightman and Thimann 1980). In addition, seaweeds con-
and S. liebmannii treatment at 1.0 %. tain precursors of elicitor compounds that promote germina-
In tomato, high concentrations of salt in the germination tion (Stephenson 1974), growth, and maintenance of plant
media significantly delayed the onset, reduced the rate of health (Kloareg et al. 1996). Another possibility is the pres-
germination (Foolad and Lin 1997, 1998), and reduced the ence of polysaccharides in LSEs, as sugars that are known to
germination percentage (Hajer et al. 2006). In this study, the improve plant growth in a similar way to hormones (Rolland
reduction in germination and growth (plumule length and rad- et al. 2002). Zeatin is another candidate for induction of
icle length) following applications of LSEs of C. sertularioides rooting in plants by seaweed (Finnie and van Staden 1985).
and S. liebmannii could be a result of high-salinity extracts. Furthermore, brown and green seaweed extracts contain
Seed germination of tomato lines (LA3770, R205, CT6, FLA, various betaines and betaine-like compounds (Blunden et al.
and ME) was delayed by salinity increasing from 2.5 to 1986; Ghoul et al. 1995). In plants, betaines serve as a com-
10 dS m−1 (Kaveh et al. 2011). In addition, germination rate patible solute that alleviates osmotic stress induced by salinity
decreased in tomato cultivars (Pascal, Red Stone, Shohba, and drought stress. However, other roles have also been
J Appl Phycol (2014) 26:619–628 625

Fig. 3 Effect of liquid seaweed Foliar spray Soil drench

extracts treatments applied as 60 a 60 b
foliar spray and soil drench on a, c c cc b bb
b shoot length, c, d root length, 50 b b 50 b b bb bb
b b
bb

Shoot length (cm)

and e, f total plant height of

Shoot length (cm)

a
40 40
tomato at different
concentrations: 0.2 % (white
30 30 aa a
columns), 0.4 % (gray columns), aa
and 1.0 % (black columns). a
20 20
Treatments: control (C), U.
lactuca (UL), C. sertularioides 10 10
(CS), P. gymnospora (PG), and
S. liebmannii (SL). Columns 0 0
C UL CS PG SL C UL CS PG SL
denoted by a different letter are
Treatments Treatments
significantly different at P≤0.05.
Values represent average (n=10
plants); bars represent standard
40 c 40 d
b
error a ab b a
a a
30 b b 30 a a
b a a a

Root length (cm)

Root length (cm)

ab a b a
a a a a
a a a
20 20

10 10

0 0
C UL CS PG SL C UL CS PG SL
Treatments Treatments

100 100
e f
90 90 c b c
80 cbb
cc 80 b bb b bb
c
Plant height (cm)
Plant height (cm)

70 b b bb 70 a
60 60 a
a a a
50 a 50
a
40 40
30 30
20 20
10 10
0 0
C UL CS PG SL C UL CS PG SL
Treatments Treatments

suggested (Blunden and Gordon 1986). It has been indicated at 1.0 % increased root length and fresh weight, whereas foliar
that betaine may work as a nitrogen source when provided in applications of the same product had no significant effect on
low concentration and serve as an osmolyte at higher concen- shoot growth but showed effect on fresh weight at a con-
trations (Naidu et al. 1987). Betaines have been shown to play centration of 0.4 %. Also, seaweed extracts enhance nutrient
a part in successful formation of somatic embryos from coty- uptake by roots (Crouch et al. 1990), resulting in root
ledonary tissues and mature seeds of tea (Akula and Bateson systems with improved water and nutrient uptake efficiency,
2000). All that information supports our results about seed thereby causing enhanced general plant growth and vigor.
germination and developed seedlings with LSEs. Our results agree with these trends. We showed that appli-
Our results relate with those in the literature (Ashok-Kumar cations of LSEs from U. lactuca and P. gymnospora as a
et al. 2012; Ganapathy Selvam et al. 2013; Kalaivanan and soil drench and foliar spray increased shoot length and root
Venkatesalu 2012; Kumari et al. 2011; Sridhar and Rengasamy length of tomato plants. In another study, Kumari et al.
2010, 2011), where seaweed extract at low concentration was (2011) reported an increase in root length, shoot length,
used for a variety of plants without any harmful effects. Effect and fresh weight in tomato treated with drench and foliar
of seaweed extract on tomato seedlings was successfully dem- applications of LSE from S. johnstonii. Enhanced vegetative
onstrated by Crouch and van Staden (1992, 1993). Tomato growth in tomato plants treated with a higher concentration
plants treated with LSE from Ecklonia maxima as a soil drench of seaweed extract could be due to the mineral nutrients
626 J Appl Phycol (2014) 26:619–628

Fig. 4 Effect of liquid seaweed a b

Foliar spray Soil drench
extracts treatments applied as
20 20
foliar spray and soil drench on a,
b fresh weight and c, d dry 18 18
c c
weight of tomato at different 16 16
b

Fresh weight (g)

bb b

Fresh weight (g)

concentrations: 0.2 % (white 14 14 b b
columns), 0.4 % (gray columns), 12 ccc 12
b
and 1.0 % (black columns). 10 bb b b 10
b bb
Treatments: control (C), U. 8 8 a
lactuca (UL), C. sertularioides 6 aaa 6 aaa
(CS), P. gymnospora (PG), and 4 4
S. liebmannii (SL). Columns 2
2
denoted by a different letter are
0 0
significantly different at P≤0.05. C UL CS PG SL C UL CS PG SL
Values represent average (n=10 Treatments Treatments
plants); bars represent standard
error c d
1.2 b b 1.2 b
b bb b
b
b b b b b bb
1 b 1 b b
Dry weight (g)

Dry weight (g)

0.8 0.8
a a
0.6 0.6
a aa aaa
0.4 0.4

0.2 0.2

0 0
C UL CS PG SL C UL CS PG SL
Treatments Treatments

present in the LSE. The beneficial effect of seaweed extract extracts helps in enzyme activation, cell elongation, and cell
application can be attributed to its many components work- stability. The organic constituents of seaweed extract include
ing synergistically at different concentrations (Fornes et al. plant hormones which elicit strong physiological responses in
2002). Similar, studies conducted with LSEs applied by low doses (Pramanick et al. 2013).
sprays under controlled experiments resulted in increased In conclusion, this study shows that liquid seaweed ex-
height and improved root growth in tomatoes (Verkleij tracts from U. lactuca and P. gymnospora were more effec-
1992; Zodape et al. 2011). Other positive results were tive at stimulating the growth of tomato seedlings, and there-
reported in spinach and tomatoes (Featonby-Smith and fore, they are potential candidates for the production of
van Staden 1983; Finnie and van Staden 1985). This en- effective biostimulants. Surprisingly, the extracts of both
hanced growth effect is thought to be due to various organ- species showed better results when they were applied at
ic compounds present in the seaweed extracts. In the pres- lower concentrations than more concentrated extracts. This
ent research, it was not possible completely establish the shows that only a small amount of seaweed extract can be
relationship between minerals and growth of tomato plants. used or even could be mixed with commercially available
However, a possible explanation for these results may be fertilizers to enhance plant growth. In places where inorganic
due to the presence of P in LSEs. The LSEs or liquid fertilizers are limited, LSEs may provide a powerful and
seaweed fertilizers can help stimulate root proliferation environmentally friendly approach to nutrient management.
and enhance root-to-shoot ratio, thereby making the plants This study provides valuable information on the identifica-
more able to mine adequate nutrients from the deeper soil tion and utilization of Mexican seaweed resources for agri-
layers and influence crop maturity as a whole. Since liquid culture. The presence of inorganic minerals in LSEs makes
seaweed fertilizer is a very good source of K, it helps in them an excellent choice as organic fertilizers. The practice
regulating the water status of the plants, controls the of applying eco-friendly seaweed extract can therefore be
opening and closing of stomata, and thereby, to a large recommended to growers to help attain better germination
extent, controls the photosynthesis. Meristematic growth, and growth of tomato or other crops. Future studies on
translocation of photosynthates, and disease resistance are specific mechanisms attributed to plant growth are required
also influenced by the presence of K. The Ca in seaweed to determine the potential of U. lactuca and P. gymnospora
J Appl Phycol (2014) 26:619–628 627

as commercial growth biostimulants which can be promoted valor nutrimental y perspectivas de aprovechamiento en la
alimentación animal. Arch Latinoam Nutr 52:115–125
as eco-friendly biofertilizers across Mexico.
Castro-González MI, Pérez-Gil R, Pérez-Estrella S, Carrillo-Domínguez
S (1996) Chemical composition of the green alga Ulva lactuca.
Acknowledgments The authors are grateful to Prof. Dr. Carla Vanessa Cienc Mar 22:205–213
Sánchez Hernández from the “Departamento de Producción Agrícola, Challen SB, Hemingway JC (1965) Growth of higher plants in response
Centro Universitario de Ciencias Biológicas y Agropecuarias (CUCBA) to feeding with seaweed extracts. In: Proceedings of the 5th
Universidad de Guadalajara” for her valuable suggestions during this International Seaweed Symposium, pp 359–367
research. Gustavo Hernández wishes to express his thanks for the fellow- Crouch IJ, van Staden J (1992) Effect of seaweed concentrate on the
ship granted under the program “Beca de exclusividad” of the “Comisión establishment and yield of greenhouse tomato plants. J Appl Phycol
de Operación y Fomento de Actividades Académicas del IPN (COFAA)” 4:291–296
and also the program “Estímulo al Desempeño de los Investigadores Crouch IJ, van Staden J (1993) Evidence for the presence of plant
(EDI) del IPN.” growth regulators in commercial seaweed products. Plant Growth
Authors thank to Biol. Kim Siewers for the English editing. Regul 13:21–29
Crouch IJ, Beckett RP, van Staden J (1990) Effect of seaweed concen-
trate on the growth and mineral nutrition of nutrient-stressed
lettuce. J Appl Phycol 2:269–272
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