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Mendel University in Brno, Czech Republic

FACULTY OF HORTICULTURE

EFFECT OF DIFFERENT MICROORGANISMS AND SUBSTRATES ON YIELD


AND FRUIT QUALITY OF CUCUMBER GROWN IN HYDROPONIC SYSTEM

DISSERATION
In partial fulfillment of the requirements of the degree of
Doctor (Ph.D.)

By
Ing. Mohamed Ewis Abdelaziz

Supervisor
Doc. Ing. Robert Pokluda, Ph.D.

Lednice December 2010

ABSTRACT
In two growing seasons of 2006 and 2007, an experiment was carried out to study the
effect of inoculation with Azotobacter chroococcum, Azospirillum brasilense, Glomus
mosseae and their combinations on growth, yield and selected component composition of two
cucumber cultivars (Cucumis sativus L. cultivars ‘Passandra’ and ‘Girola’) grown on
rockwool and peatmoss under an open soilless system in greenhouse at Faculty of
Horticulture, Mendel University, Czech Republic. Vegetative growth characteristics, number
of fruits per plant and fruits dry matter of cultivar Passandra were significantly higher than
cultivar Girola. On contrast, early and total yield did not reveal much variation between
cultivars. Nitrogen in fruits of cultivar Passandra and P, K, Ca in fruits of cultivar Girola
revealed higher significant increases in the first season while N, P, and Mg did not differ in
leaves of the two cultivars, with some exceptions. However, type of cultivar or substrate did
not affect vitamin C or TSS contents. Nitrate, recorded higher significant increase in fruits of
cultivar Girola, but values still under the critical limits. Plants grown on peatmoss revealed
significant increase in number of leaves/plant and leaf DW. Moreover, light significant
increase was obtained in early yield, fruit length and weight of cucumbers grown on
rockwool. All inoculation treatment improved total yield by 8 and 21 % in 1st and 2nd seasons,
respectively. In addition, significant increase in average fruit weight was found when plants
inoculated with Azotobacter alone or combined with Azospirillum than control in the 1st
season. Microorganisms inoculation improved the marketable yield, mineral component in
leaves and fruits of cucumbers, with some exceptions. Vitamin C recorded the highest
significant increase with Glomus + Azospirillum. Furthermore, N-Fixers alone/or combined
with Glomus improved growth of cucumber plants grown in peatmoss. In particular, growth
and yield of cultivar Passsandra grown in peatmoss were promising responded to
microorganisms inoculation than other treatments under the open soilless culture.

Keywords: cucumber, soilless culture, peatmoss, rockwool, Azotobacter chroococcum,


Azospirillum brasilense, Glomus mosseae, growth, yield, nutritional value
ABSTRAKT

V letech 2006 a 2007 byly provedeny experimenty studující vliv inokulace Azotobacter
chroococcum, Azospirillum brasilense, Glomus mosseae a jejich kombinací na růst, výnos a
vybrané obsahové složky okurek Cucumis sativus L. (odrůdy ‘Passandra’ a ‘Girola’)
pěstovaných v čedičové plsti a v rašelině v otevřeném hydroponickém systému ve skleníku
Zahradnické fakulty v Lednici, Mendelovy Univerzity v Brně, Česká Republika.
Vegetativní růstové charakteristiky, počet plodů na rostlinu a obsah sušiny plodů byl
průkazně vyšší u odrůdy Passandra. Na druhé straně ranost a celkový výnos nebyl odrůdou
ovlivněn. Obsah dusíku v plodech odrůdy Passandra a obsah P, K, Ca v plodech odrůdy
Girola byl průkazně vyšší v první sezóně. Na druhé straně obsah N, P a Mg v listech byl mezi
odrůdami, až na několik výjimek, srovnatelný. Podobně genotyp neovlivnil obsah vitaminu C
ani TSS. Obsah dusičnanů byl vyšší u odrůdy Girola, ale stále pod hygienickým limitem.
Rostliny pěstované v rašelinovém substrátu vykazovaly vyšší počet listů na rostlinu a vyšší
zastoupení sušiny v listech oproti variantě v čedičové plsti. Ovšem čedičová plsť přispěla
k průkaznému, byť mírnému, zlepšení ranosti, délky a hmotnosti plodů okurek.
Všechny inokulované varianty vykázaly vyšší celkový výnos o 8 a 21 % v 1. a 2. pokusném
roce. Také byl zjištěn průkazný nárůst hmotnosti plodů při inokulaci rostlin Azotobacter
samotným nebo v kombinaci s Azospirillum, ve srovnání s kontrolou. Inokulace také zvýšila
tržní podíl výnosu, zastoupení minerálních látek v listech a plodech okurek, i když s několika
výjimkami. Obsah vitaminu C byl průkazně zvýšen ve variantě ošetřené Glomus +
Azospirillum. Vedle toho N-fixujíxí bakterie samostatně nebo v kombinaci s houbou Glomus
podpořily růst okurek pěstovaných v rašelinovém substrátu. Obzvláště růst a výnos odrůdy
Passandra v rašelinovém substrátu ukázal pozitivní odpověď na inokulaci ve srovnání
s dalšími sledovanými variantami v otevřené hydroponické kultuře.

Klíčová slova: okurky, hydroponie, rašelina, čedičová plsť, Azotobacter chroococcum,


Azospirillum brasilense, Glomus mosseae, growth, yield, nutriční hodnota
CONTENTS

Page

ABSTRACT ii
I. INTRODUCTION 1

II. REWIEW OF LITERATURE 3


II .1. Crop 4
II. 2. Effect of substrate 6
2.1. Type of substrate 9
2.1.1. Rockwool 10
2.1.2. Peatmoss 11
2.2. Vegetative growth characteristics 14
2.3. Yield and its component 15
2.4. Chemical components 17
II .3. Effect of microorganisms 19
3.1. Nitrogen fixing bacteria 19
3.2. Arbuscular mycorrhiza 23
3.3. Vegetative growth characteristics 26
3.4. Yield and its component 29
3.5. Chemical components 30

III. MATERIALS AND METHODS 32

IV. RESULTS AND DISCUSSION 38

IV.1. Vegetative growth characteristics 38


1.1. Effect of cultivar 38
1.2. Effect of substrate 38
1.3. Effect of microorganisms inoculation 41
1.4. Effect of interactions 44

IV. 2. Yield and its component 49


IV.2.1. Early and total yield 49
2.1.1. Effect of cultivar 49
2.1.2. Effect of substrate 49
2.1.3. Effect of microorganisms inoculation 51
2.1.4. Effect of interactions 53
IV.2.2. Yield component 56
2.2.1. Effect of cultivar 56
2.2.2. Effect of substrate 56
2.2.3. Effect of microorganisms inoculation 58
2.2.4. Effect of interactions 60
IV.2.3. Marketable yield 63
2.3.1. Effect of cultivar 63
2.3.2. Effect of substrate 65
2.3.3. Effect of microorganisms inoculation 66
2.3.4. Effect of interactions 66

IV. 3. Mineral components 71

IV. 3.1. Mineral composition of leaves 71


3.1.1. Effect of cultivar 71
3.1.2. Effect of substrate 71
3.1.3. Effect of microorganisms inoculation 73
3.1.4. Effect of interactions 75

IV. 3.2. Mineral composition of fruits 82


3.2.1. Effect of cultivar 82
3.2.2. Effect of substrate 82
3.2.3. Effect of microorganisms inoculation 84
3.2.4. Effect of interactions 86

IV. 4. Chemical quality 91

IV. 4.1. Vitamin C and total soluble solid (TSS) 91


4.1.1. Effect of cultivar 91
4.1.2. Effect of substrate 91
4.1.3. Effect of microorganisms inoculation 91
4.1.4. Effect of interactions 94

IV. 4.2. Dry matter content, nitrate and total sugars concentrations 98
4.2.1. Effect of cultivar 98
4.2.2. Effect of substrate 98
4.2.3. Effect of microorganisms inoculation 100
4.2.4. Effect of interactions 102
V. SUMMARY 104
VI. REFERENCES 110
VII. APPENDIX 126
ACKNOWLEDGEMENT
First of all I would like to present this work to Ing. Samah Shaban to express my
deep feeling towards her support and help to finish this work.
The author is deeply grateful to Associate Prof. Robert Pokluda, Head of
Vegetable Growing and Floriculture Department for his ideal supervision, willing
advice, continuous encouragement, valuable help during the course of this investigation.
I am also glad to extent my sincere gratitude to Associate Prof. Kristina
Petříková, CSc. for sharing constructive ideas, necessary support and required
information. I do appreciate and thank Mrs. Jarmila Turečková (Secretary of the
Department) for her persistent help.
Finally, I would like to extend my deepest appreciation for the Czech Ministry of
Education, Youth and Sports (Czech Republic Foreign Aid Programme), the
Government of Czech Republic for the acknowledged scholarship grant and enabling
me to join and complete an intended further study.

Lednice, December 2010


I. INTRODUCTION
Cucumber (Cucumis sativus L.) is one of the most important vegetable crops in the
world grown under greenhouses.
High yield and good quality under open field condition need extreme care of soil
fertility and soil handling, specially using of organic manure and deep soil preparation
(Peyvast et al, 2008). On the other hand, soilless culture allows a more accurate control of
root environment which offers possibilities for increasing production and improving quality
(Bohme et al, 2001).
Substrate culture is the main popular hydroponic system between the various
hydroponic techniques (Maloupa et al, 2001). In Europe, there are now 6000 ha of
hydroponics and an expected area of 10000 ha in the coming years (Van Os, 2001).
Recently, many materials are used as a growing media such as rockwool, pumice
stone, expanded clay and perlite (Schie, 1999). Using of different organic and inorganic
substrates allow better nutrient uptake, sufficient growth and development due to optimizing
water and oxygen holding (Verdonck and Demeyer 2004).
Most of the greenhouse farms in Europe with the hydroponical substrate culture are
using rockwool slabs (Bohme et al, 2008). Many of them have problems with the reuse and
recycling of rockwool in the greenhouse. The use of organic and completely compostable
substrates is therefore of rising interest in hydroponics. In this respect, slabs of peat have been
used successfully and with increasing amount (Bohme and Vorwerk 2003).
Recently, there has been a lot of work conducted to improve protected culture
practices in European countries. These practices particularly involve application of fertilizers
in excess to insure that there is no shortage in the nutrient solution through different stages of
plant life.
In commercial application, type of substrates and mineral fertilization are very
important factors affecting the production and quality of vegetables under soilless culture
conditions, especially the balance between N, P and K in growing media and/or nutrient
solution.
Generally, open soilless culture systems have been associated with pollution problems
which allow drain out the excess nutrient solution. In connection, the recent environmental
regulations against ground water and soil pollution restrict using of open system.
Moreover, increasing the applications of NPK fertilizers does not sustain economical
production and cause many problems for environment and human health, especially with
increasing levels of NO3- (Hanafy Ahmed et al, 2004). Therefore, it is a place of important to
nourish plants by natural sources for macro nutrients, especially N, P and K, which could
increase plant production without contaminating environment.
Azospirillum and Azotobacter are plant growth promoting rhizobacteria (PGPR)
capable of affecting the growth and yield of numerous plant species, cause many of
agronomic and ecological significant effects (Bashan and Holguin, 1997; Bashan et al, 2004;
del Amor et al, 2008). In addition, arbuscular mycorrhiza (AM) is involved in the most
durable, intimate and important symbiosis on the earth (Allen, 1996; Hodge, 2000), and form
mutualistic symbioses with a vast majority of land plants (Smith and Read, 1997).
PGPR not only have been reported to enhance plant growth directly by a fixation of
atmospheric nitrogen that is transferred to the plant, but also it have the ability to produce
siderophores that chelate iron and make it available to the plant root, solubilization of
minerals such as phosphorus, and synthesis of phytohormones (Glick, 1995; Dobbelaere et al,
2001).
The aim of this work was to study the effect of inoculation with Azotobacter,
Azospirillum and Glomus on growth, yield and selected component composition in leaves and
fruits of cucumber plants grown on rockwool and peatmoss as based growing media in an
open soilless system.
II. REVIEW OF LITERATURE
Nowadays, there is an urgent need to sustain vegetable cultural practices on global
level, particularly under soilless conditions. In the developed world a reduction of energy and
environmental costs is necessary. In developing countries, efficient and sustainable practices
are needed to allow cost-efficient production of adequate nutrition for the growing
populations. To overcome the ecological problems and to increase crop yields, microscopic
organisms that allow more efficient nutrient use for or increase nutrient availability can
provide sustainable solution for present and future agricultural practices (Rai, 2005).

A stable substrate “growing medium” is essential for successful growing of range of


species, basis on the small rooting volume for vegetable crops.

Thus, water uptake, buffering and availability as well as oxygen diffusion must be
optimal. Also pH and concentration of nutrients and other salts should meet the plant
demands as well as biological activity should be such kind does complete with the plants
(Verhagen, 2009). Therefore, the bellow factors will be reviewed:

1. Crop

2. Substrate “growing media”

3. Microorganisms inoculation

In addition, the effect of microorganisms inoculation and type of substrate on various


vegetable plants will be discussed according to their effects on:
1. Vegetative growth characteristics

2. Yield and its component

3. Mineral components

i. Mineral composition of leaves

ii. Mineral composition of fruits

iii. Chemical quality

II. 1. Crop
Cucumbers (Cucumis sativus) belong to the same family, Cucurbitaceae, as
watermelon, zucchini, pumpkin, and other types of squash. Greenhouse cucumber production
is very popular in many areas of the world. The cucumber is a warm season crop with
required growing conditions of 27 to 30 oC and plenty of sunlight.

Most of European greenhouse cultivars are parthenocarpic. The European seedless


type of cucumber became widely grown in many areas. The fruits are mild in flavor, seedless,
and have a thin edible skin that requires no peeling. Harvested fruit generally range from 300
mm in length and weigh about 450 g each. However, smaller fruits of mini cucumbers have
become more popular and its commercial production has increased.

Cucumber plants are indeterminate in growth, continually producing fruit on new


growth, similar to greenhouse tomatoes. Greenhouse cucumbers are more sensitive to low
temperatures than tomatoes. Minimum temperatures should not be lower than 18 °C for
sustained production. Prolonged temperatures above 35 °C should also be avoided as fruit
production and quality are reduced at extremely high temperatures.

A fruit may be developed at each node and more than one fruit may begin to develop
at some nodes. It is usually best to thin these multiple-fruit clusters to a single fruit; however,
vigorous plants can sometimes mature more than one cucumber at a node. Miniature and beit-
alpha types will support several fruit per node. Any distorted fruit should be removed
immediately. The greatest growth of the fruit occurs between day 6 and 14 after the bloom
opens (anthesis). Maximum fruit length occurs at day 14 followed by diameter increase.
(Hochmuth, 1990)

Cultivars of cucumber are grown either to be eaten fresh or to be pickled. Those that
are to be eaten fresh are commonly called slicing cucumbers. Cucumbers such as gherkins
that are specially cultivated to make pickles are oftentimes much smaller than slicing
cucumbers.

Accurate statistics on the acreage of greenhouse devoted cucumber production are not
easily obtained as no official accounting is made by any government or private organizations
(Benton, 2005).

Significant advances continue to be made in the application of hydroponic/soilless


culture methods of growing and will be continue to be mad for controlling the environment
within the greenhouse as well as the introduction of plant cultivars better adapted to
greenhouse conditions.
Table 1. Some nutritional values of cucumbers,
slices, with peel 100 g.*
Nutrient Amount
Energy 52 kJ
Dietary fiber 0.8 g
Carbohydrate 2.16 g
Folate 14 mg
Calcium 14 mg
Iron 0.22 mg
Magnesium 12 mg
Manganese 0.073 mg
Phosphorus 21 mg
Potassium 136 mg
Protein 0.59 g
Sugars 1.38 g
Vitamin C 3.2 mg
* ( www. 1)
II. 2. Effect of substrate
Growing media are commonly evaluated with respect to their physical (bulk density,
porosity, water retention capacity, air capacity) and chemical properties (readily available
nutrients, cation exchange capacity, salinity, pH and load of bulk and pollutants) (Alsanius
and Wohanka, 2009).

Quality of growing media is largely defined as the ability to perform various intrinsic
and extrinsic functions, presented by a suite physical, chemical and biological prosperities,
namely to:

(i) Provide a medium for plant growth and biological activity

(ii) Regulate and partition water flow and storage in the environment

(iii) Serve as an environmental buffer in the formation and degradation of


environmentally hazardous compounds

Carbon turnover is a central factor for biological qualities of growing media. In


accordance, Figure (1) outlines the concept of carbon turn over in growing media to elucidate
major events during the use of growing media that affect biological quality. During the life
time of a growing media, there is a steady turnover in carbon. As Carlile and Wilson (1991)
mentioned that, upon adequate storage, inorganic growing media may almost be sterile before
the first use.

In contrast, the composition of organic growing media, their C/N ratio and
environmental conditions during storage is integral to the growing media biological qualities.
Bohme et al (2008) mentioned that organic substrates show a higher CO2 outflow than
mineral medium.

The environmental conditions for the microorganisms inhabiting the growing media
are continuously changing, as the microbial density increases and the substrate concentration
simultaneously decreases.

The most dramatic change, however, occurs with the introduction of the plant or the
transplant stage, as a producer of organic carbon is introduced into the system. Root exudates
release a pool of organic carbon (Whipps, 1990) which in turn affects the composition of the
micro flora and microbial mediated processes.
Fig. 1. Major events during the use of growing media in terms of carbon turn over*

GM production site

Introduction
of the plant Treatment
prior to reuse Introduction
of the plant

Storage First use Reuse

Statistic view Dynamic view


*Alsanius and Wohanka (2009).
Furthermore, plant stress, dilution rate and the concentration of the limiting nutrients
(carbon, nitrogen and micronutrients) are important factors. At very low concentrations, i.e. in
the beginning of a crop grown in mineral growing media, the microbial rate is reduced.
Nutrients are not transported fast enough into the cell in order to match its metabolic
demands. Under moderate or high nutrient concentrations, in a crop grown in organic growing
media or in a late stage of the crop, the microbial growth rate may not be affected; meanwhile
the cell density may increase (Alsanius and Wohanka, 2009).

The influx of root or growing media originating organic carbon and the added water
volume have sustainable impact in the horticultural system. If irrigation is performed at a high
drainage percentage, microbial growth is limited and microbial cells may even be washed out.

In contrast, low dilution rates occur when the added volume water corresponds to the
plant’s water demand or when the carbon influx is low. The first case will lead to an increase
in cell density where as the latter one may cause cell death due to starvation, as limiting
nutrients may not be available fast enough to permit maintenance of cell metabolism. If the
growing media with a plant is considered from a strictly microbiological point of view,
substrate limited growth can be maintained as long as essential nutrient factor, such as H+
donors, C, N, S or P sources, are available. As long as the actual growth constant (Fuchs,
2006) is lower than maximum at substrate saturated conditions, the dilution rate may be
varied over a large range without washing out microorganisms.
Different investigations show a stability in microbial density over time (Berkelmann,
1992); however, shifts occur with respect to the microbial community structure (Khalil and
Alsanius, 2001; Bergstrand et al, 2009).

Moreover, Wallach et al (1992), Da Silva et al (1993) and Naasz et al (2005) showed


that hydraulic conductivity of various horticultural substrates (peat, composed bark,
agricultural waste compost, etc.) radically decreases on restricted ranges of water potential
that could rapidly lead to very weak water flows in the substrate and that are potentially
limiting for the root environment.

Many years ago, Puustjarvi (1997) described a 20-25% decrease of peat volume in
carnation culture. The rapid and extreme variations in water and air availability to roots lead
to frequent cycles of watering (fertigation) and drying (evaporation, root uptake) during
growing management called shrink/swell phenomena (Heiskanen, 1995 and Michel et al,
2004).
Table 2. Bulk density of the substrates used in cucumber production (g.L-1)*

Stages
Media type After After
New
first use second use+
Perlite
132 151 106
Coconut fiber
124 168 176
Rockwool
50 70 106
Peat slabs
133 171 177

* +
Bohme, et al (2008) with dried roots

This phenomena lead to a change in solid phase and pore organization, which could
then influence the water and air circulation in growing media (Table 2), leading to a decrease
in aeration that could cut off the supply of oxygen to the root during growth (Lemaire et al,
2003).

High irrigation frequency may improve crop performance due to a higher availability
of nutrients, especially P and Mn, while maintaining constantly higher moisture levels in the
root zone.

The only precaution regarding the application of a frequent irrigation schedule is the
possible creation of excessive moisture condition in the root zone that might reduce oxygen
availability (Savvas, 2009). Poor oxygen diffusion leads to lack of oxygen which causes die
off roots (Verhagen, 2009).

In addition, peat breakdown could lead to a more open structure with higher air filled
porosity. Weakly decomposed peat has an open structure which is sensitive to breakdown. In
practice is seen that peat materials with a high bulk density are less sensitive than those with
low bulk densities. Cattivello et al (1997) reported that particle size it self is not an important
factor in shrinkage of peat materials. On the other side, Verdonck and Demeyer (2004)
revealed that particle size is an important factor in determine the water and air relationship in
peat substrate.

These interactions show, that physical and biological proprieties of growing media
cannot be considered independent from cultural measurers directly (e.g. fertilization) or
indirectly (e.g. affecting carbon partitioning with the plant) the plant nutritional situation in
the root zone. In this context it has to be pointed out that microbial communities associated
with roots and growing media are extremely complex.

2.1. Type of substrates


Soilless culture allows a more accurate control of root environment which offers
possibilities for increasing production and improving quality (Van Winden, 1988). The
interest in commercial application of soilless culture has been increasing since it is an
excellent alternative to soil disinfection and increase the water use efficiency (Gul et al,
2009).

The most common growing media for greenhouse vegetable production are rockwool
(Tayson et al, 2001), perlite and gravel (Hamdy et al, 2002) as mineral medias. Currently,
more than 50 % of greenhouse vegetables are produced using rockwool as a growing media
(Zheng et al, 2009).

However, several factors must be considered when selecting a media for soilless
hydroponic vegetable production. A media must drain freely, especially as it ages, be easily
aerated, have good water holding capacity, be nontoxic, and free of weeds, pests, disease and
chemicals (Cantliffe et al, 2003). Furthermore, the media must be inexpensive (Cantliffe et al,
2001).

2.1.1. Rockwool
Rockwool is the most widely used inert substrate in soilless systems (Martyn, 1996;
Bohme et al, 2006) and it has been considered the most suitable medium to utilize the
following hydroponics-characteristics:

(i) Chemically inactive and the nutrient solution is not affected

(ii) May hold a large amount of solution and has optimal aeration

Therefore, optimal moisture distribution is maintained in the medium, which enhances


root development and formation of the root system (Smith, 1987; Tanimoto, 2002) and
provide sufficient levels of oxygen to the roots (Raviv et al, 2002). In addition, rockwool is
more lightweight when dry, easily handled and steam-sterilized than many other types of
materials (Inden and Torres, 2004).

Table 3. Physical properties of mineral media used in greenhouse culture*

Physical property
Media type Air
Moisture
porosity pH
capacity (% vol.)
(%)
Coarse Perlite
17 40 6.9
Rockwool
80 18 7
*
Prasad (1997)
On contrast, it can’t be denied that using of rockwool slabs has also many
disadvantages such as:

(i) There is a substantial loss of unused water and nutrient elements when the
nutrient solution flows from the slabs

(ii) Root growth may become as massive as to fill the slab leading to root death (O2
starvation) and root may grow out of the drain holes in the base of the sleeve
enclosing the slab

(iii) The slab requires periodic leaching to remove accumulated salts is order to
prevent a soluble salt buildup, a determination based an electrical conductivity
(EC) measurement made on a draw liquid sample from the slabs or effluent from
the slabs (Benton, 2005).
Fig. 2. Porosity of inorganic materials (a) volcanic scoria, (b) perlite and (c) pumice.
(Anicua-Sanchez et al, 2008)

(a) (b)

(c)

Increased environmental and economic pressure pushes the horticultural industry to


find substitute substrates for the rockwool slabs used in greenhouse production (Allaire et al,
2004).

2.1.2. Peatmoss
Some growers use non-mineral media such as pine bark, coconut coir and sphagnum
peat mixtures because of their satisfactory aeration and water holding capacity (Cantliffe et al,
2003; Iapichino e al, 2007).

Peat has been the main raw material for industrially produced growing media, in
particular in greenhouse production (Sarkka et al, 2004). The availability of peat in large
quantities of relatively high and homogenous quality is necessary for different purpose. The
high percentage of peat in substrates makes the quality aspects very important (Keijzer,
2003).
Table 4. Physical properties of organic media used in greenhouse culture*

Physical property
Media type Moisture
Air porosity (%) pH
capacity (% vol.)
Coir
33 16 5
Pine bark
17 46 5.2
Sphagnum peat
50 12 5.8
*Cantliffe et al (2003)

Fig. 3. Porosity of organic materials (a) coconut fiber, (b) sphagnum peat and (c)
vermicompost. (Anicua-Sanchez et al, 2008)

(a) (b)

(c)

In this respect, Verdonck and Demeyer (2004) declared that the physical of growing
media become more and more important for optimal plant growth. As they can not be
changed during the growing period, these properties should be optimal from the beginning. In
contrast, the chemical properties can be adapted to optimal conditions during the growth
period. The authors mentioned that particle size is an important in determine the water and air
relationship in peat substrate, as shown in the Table (5).
Table. 5. Physical properties of different fraction of peat, bark and perlite*

Physical property
Media type Volume Volume
% air % EWA+
Peat 0-1 mm
6.9 35.5
Peat 1-2 mm
37.5 27.4
Peat 2-3 mm
50.4 13.1
Composted bark 0-1 mm
12.6 24.6
Composted bark 1-2 mm
38.2 9.0
Composted bark 2-3 mm
53.6 3.1
Perlite 0-1 mm
20.4 42.9
Perlite 1-2 mm
57.7 1.2
+
*Verdonck and Demeyer (2004) Easy available water
Due to very large peat reserves, light sphagnum peat has been the main raw material
for horticultural use, while darker decomposed peats have been used mainly for energy
production. However, when using growing media mainly containing light peat, problems may
arise, especially in long term use.

Light sphagnum peat possesses a range of positive and characteristics, including low
bulk density, good water holding capacity.

On the other side, the difficulty of obtaining positive impacts of AM on plant growth
in peat based growing media has been recognized for some time (Estaun et al, 1994).

There are several possibilities for this like the excess amount of P in the substrate, the
low anion holding capacities of sphagnum peat lead to increase the need for frequent fertilizer
application in greenhouse culture, and AM suppressive microorganisms in peat may be
another possible reason for weak AM symbiosis in peat based growing media. Also, peat
growing media have high populations of fungi and bacteria, in particular actinomycetes,
which may affect symbiosis efficiency of AM (Vestberg and Kukkonen, 2009). In addition,
extra cellular enzymes like ß-glycosidase, phosphatase, sulphatase, xylosidase, esterase
present in peat (Freeman et al, 1995) may interact with AM.

Linderman and Davis (2003) demonstrated that different peat substrates indeed have
some deleterious effect on mycorrhiza formation and the potential for mycorrhiza induced
plant growth enhancement under P limiting conditions.

The authors concluded that the nature of the inhibitory factor, however, remains
unknown. This factor may be found in the organic chemistry of peat. Peat consists of mainly
humines, humic and folic acids, cellulose and hemi cellulose, bitumen, pectin and small
amount of mineral substances. Some of this component may interact with AM.

However, this effect can vary also with AM strain (Corkidi et al, 2004). In contrast,
humic acids have shown a positive impact on AM development and function (Gryndler et al,
2005).

Certain peat qualities can even possess properties that seem to have a negative effect
on AM symbiosis. Incompatibility between AM and peat, the reasons of which are still
largely unknown, is maybe the largest obstacle for using AM in peat-based growing media
(Vestberg et al, 2000; Linderman and Davis, 2003).

Slabs of peat and coconut fibers have been used successfully and with increasing a
mount (Bohme and Vorwerk, 2003). Tahvonen and Kemppainen (2008) reported that light
sphagnum peat has proved to be the best and most reliable substrates for plants in
greenhouses when aiming for higher yields.

However, poor understanding of substrates leads to production difficulties and the


inefficient use of water and nutrients (Parks et al, 2009 a).

So that, the effect of different substrates will be reviewed as follows in the next
chapters:

2.2. Vegetative growth characteristics


Mohamed and Enzy (2001) found that plant height of sweet pepper plants were
increased significantly when plants grown on sandy soil, sand bags and gravel systems when
compared with NFT system. In addition, Peyvast et al (2008) mentioned that plant fresh
weight was significantly higher in cucumber grown in peat than perlite substrates.
In connection, Shahinrokhsar and Tavusi (2008) revealed that number of leaves, fresh
and dry weight of strawberry plants grown in coco peat were significantly higher if compared
to those grown in bark + perlite (1:1 W) substrate.

On the other hand, Cantliffe et al (2003) declared that type of media (perlite, coarse
perlite, and pine bark) did not affect stem length in pepper plants. Similarly, Shinohara et al
(1999) found no differences in growth characters between tomato plants grown in coconut
fiber or rockwool.

In cucumber, number of leaves, stem diameter as well as plant height in plants grown
in pumice or perlite did show any differences among the substrates (Abul-Soud et al, 2003).

Recently, AL-Rawahy et al (2009) did not find differences between the two substrates
(wood straw and date palm leaf straw) in terms of plant height and stem thickness for
cucumber plants grown in open soilless system. Also, Parks et al (2009a) revealed no
difference between medium contained: equal share of sand, perlit, composted pine bark and
coir, by volume, when compared to another medium contained 25 % sand, 25 % perlite, 50 %
composted pine bark in terms of cucumber shoot dry weights.

However, Inden and Torres (2004) declared that fresh and dry weights of tomato were
significantly higher in plants grown in perlite plus carbonized rice hull, while no differences
in fresh and dry weights of the 4th leaf were recorded between plants grown in rockwool or
coconut coir substrates. In addition, Ikiz et al (2009) studded the effect of three different
application treatments of mycorrhiza on pepper plant grown in soilless culture. The authors
found that dry weight of the plants grown in perlite medium had 28 % increases by one time
mycorrhiza application and 41 % increases by two time mycorrhiza application if compared
to control.

2.3. Yield and its component


The soilless cultivation is one of the most efficient alternatives to soil fumigation since
it provides a pathogen-free root environment at planting, while promoting earlier and higher
yields (Savvas, 2009).

However, effects of different substrates on crop yield and its component are varying
among grown crops. In this respect, Maniutiu et al (1995) found that cucumbers plants grown
in bags of peat showed significant increase in early and total yield compared to the plants
grown in straw beds.
Moreover, Lee BeomSean et al (1999) pointed out that greater marketable yield and a
lower incidence of abnormal fruits were observed on a substrate of perlite with rice hull or
with carbonized rice hull as compared to the pure perlite substrate. In addition, Broeck
(1999), declared that cucumber plants cultivated in pumice produced the highest fruit number,
with slightly lower average crop yield than rockwool substrate.

Mohamed and Enzy (2001) found that early yield of sweet pepper plants was
increased significantly when plants grown on sandy soil, sand bags and gravel systems when
compared with NFT system. Moreover, Tuzel et al (2001) mentioned that tomato grown in
perlite + peat resulted higher fruit number per plant than plants grown in perlite alone.

In connection, Colla et al (2003) reported that marketable yield per plant was
significantly higher in cucumber plants grown in perlite and coir than those grown in
rockwool. The authors also mentioned that high fruit number and average fruit weight of
plants grown in perlite and coir caused these significant difference compared to plants grown
in rockwool. These results were confirmed by Parra et al (2009) on tomato, which found that
marketable yield for tomato grown in coir was significantly higher than those grown in
rockwool under open system condition.

Moreover, Peyvast et al (2008) pointed out that total marketable yield (9.0 and 7.9
kg.m-1) and numbers of cucumber fruits (102 and 92 m2) were significantly higher with plants
grown in peat than perlite, respectively. The authors found that peat substrate produced the
highest fruit fresh weight with 9.8 kg.m2 followed by perlite and perlite + rice hull with an
amount of 8.6 and 8.1 kg.m2, respectively. Meanwhile, different substrates had no significant
effect on the non-marketable yield. Similar trend was obtained by Show et al (2004) on ‘Beit
Alpha’ cucumbers.

Recently, Parks et al (2009 b) mentioned that mean fruit weight was approximately
48 % higher for cucumber and 12 % higher for tomato grown in coir compared to the other
substrates (three component including sugar cane waste, peanut husks, soybean, peat or
volcanic rock).

On the other hand, the highest tomato fruit yield was obtained for plants grown in
rockwool, compared with plants grown in expanded clay or polyurethane foam (Chohura and
Komosa, 2000) or coconut fibers (Benko et al, 2008). In addition, Bohme et al (2008) found
that yield was very low in cucumber plants grown in peat slabs compared to those grown in
rockwool. The authors explained this increase to the rockwool ability to absorb more nutrients
than peat slabs.

Meanwhile, many authors found that yield of cherry tomato cultivated on coconut
fiber was similar to the yield on rockwool (Holtmann and Kobryn, 2003; Inden and Torres,
2004). Lopez-Galarza et al (2005) pointed out that type of substrate (coir dust, perlite and
rockwool) did not affect yield of watermelon plants

In this respect, Cantliffe et al (2003) declared that type of media (medium perlite,
coarse perlite, and pine bark) had no significant effect on fruit yield of cv. Beit Alpha
cucumbers (Show et al, 2004) as well as fruit set and total number of nodes per plant in
peppers. While, in strawberry, plants grown in perlite produced a higher marketable fruit
number and fruit weight per plant compared to pine bark and mixture of 2 peat: 1 perlite.

Also, Schahinrokhsar and Tavusi (2008) found no significant differences in yield, fruit
number and fruit weight of strawberry grown in coco peat or bark substrates.

In cucumber, Schroeder and Sell (2009) found no significant differences in yield for
‘Euphoria’ and ‘Aviance’ cultivars grown in peat or composted substrate.

Recently, no difference in fruit number and fresh weight of cucumbers and tomatoes
were found between medium contained 25 % sand, 25 % perlite, 25 % composted pine bark
and 25 % coir by volume when compared to medium contained 25 % sand, 25% perlit, 50%
composted pine bark (Parks et al, 2009 a).

2.4. Chemical component


Plant nutrition and fruit yield from different cultivars of cucumber under different crop
systems, have been studied in the literature (Gomez et al, 2002 a), but the corresponding
systematic studies about differences in cucumber quality throughout the harvest season
(Schouten et al, 2002), or indifferent stages of fruit development are not common for different
production systems (Fernandez-Trujillo et al, 2004).

Tuzel et al (2001) found that fruit of tomato showed significant increases in dry
matter, vitamin C and total soluble solid content when grown in volcanic ash compared to
perlite substrates. Also, fruits of cherry tomato grown on coco-fiber contained much more
total sugars, while those grown on rockwool decidedly in vitamin C (Holtmann and Kobryn
2003). Similarly, Inden and Torres (2004) found that the high soluble solids content was
higher in plants grown in coconut coir (7.3) followed by rockwool (6.6O Brix) and perlite
(5.6O Brix).

In addition, Parks et al, (2009 b) studied the effect of four substrates on quality of
cucumber fruits. The authors found that nitrate concentration in cucumber was particularly
higher for sugar can waste, peanut husks and peat medium, while vitamin C recorded higher
content in the coir medium.

On the other hand, Guler et al (1995) did not find differences in soluble solid content
among melon fruits grown in rockwool, perlite, coir slabs or coir dust + perlite mix. Also,
quality of cucumbers grown in straw beds or bags of peat substrates (Maniutiu et al, 1995) did
not show any differences among substrates. Similar results were obtained with tomato grown
in coconut fiber and rockwool (Shinohara et al, 1999).

Furthermore, Colla et al (2003) revealed no significant differences in fruit quality


between cucumbers grown in rockwool, perlite and coir. Moreover, the author mentioned that
dry weights of fruits were higher in plants grown in coir and perlite than those grown in
rockwool. Similarly, Peyvast et al (2008) found no significant effects on fruit length, diameter
and soluble solid in cucumber grown in peat, perlite or rice-hull substrates. Fernandez-Trujillo
et al (2004) reported that differences in soluble solid of cucumber were mainly affected by
harvest date than to the type of growing systems (NFT or perlite substrate). The authors
mentioned that titratable acidity during the winter season was overall 14 mmol H+ L-1, but
during the spring season remained higher in perlite grown fruit than NFT.

Also, strawberry plants grown in coco peat and bark substrates did not reveal
significant differences between the two substrates (Schahinrokhsar and Tavusi, 2008). Al-
Ajmi et al (2009) found that K and P concentrations were higher in fruits of cherry tomato
grown in zeolite if compared to perlite substrate, while fibbers and protein contents were not
affected by substrates.

According to macro and micronutrients, Colla et al (2003) investigated the effect of


different substrates on mineral composition of cucumber grown in soilless culture. The
authors found that mineral composition of nutrient solution was significantly affected by type
of substrate. Nitrogen uptake is considerably higher in cucumber grown in NFT systems if
compared to perlite substrate. Lower but still significant differences were obtained for K, Ca
and Mg, while the uptake of P was not significantly affected (Gomez et al, 2002 b).
Potassium, phosphorus and sodium concentration was higher in coir than rockwool
while magnesium concentration was not significantly affected by the type of substrate. In
contrast, Schroeder and Sell (2009) mentioned that nutrient content of cucumber fruits grown
in peat or compost did not differ significantly between the substrates.

II. 3. Effect of Microorganisms


During the past four decades we have witnessed the doubling of the human population
and food production (Vance, 2001). Plant nutrition has played a key role in this dramatic
increase in demand for and supply of food (Rai, 2005). Increases in crop production have
been made possible through the use of commercial fertilizers. The use of nitrogen has almost
increased nine fold and phosphorus more than four fold (Vance, 2001). The increasing use of
fertilizers and highly productive systems have been created environmental problems such as
deterioration of soil quality, surface water, and groundwater, as well as air pollution, reduced
biodiversity, and suppressed ecosystem function (Socolow, 1999; Vance, 2001).

3.1. Nitrogen fixing bacteria


The most limiting nutrients for plant growth are N and P (Schachtman et al, 1998).
Most of N is tied into the soil organic matter. Even after fertilization, plants have to compete
with soil microbes for easily available soluble N. Problems with P are different. In acidic
soils, even when added substantial quantities as fertilizers, P precipitates with iron or
aluminum, whereas in alkaline soils P precipitates as calcium phosphates (Hinsinger, 2001).

Atmospheric N2 composes approximately 80 % of the air we breathe. Although


abundant and ubiquitous in the air, N is the most limiting nutrient to plant growth because the
atmospheric N is not available for plant uptake. It is a molecule composed of two atoms of
nitrogen linked by a very strong triple bond. Large amounts of energy are required to break
this bond and the molecule is therefore quite chemically nonreactive. The general chemical
reaction for the fixation of nitrogen (N + 3H2 + Energy -> 2NH3) is identical for both the
chemical and the biological processes. Living organisms use energy derived from the
oxidation ("burning") of carbohydrates to reduce molecular nitrogen (N2) to ammonia (NH3).
The triple bond of N must be broken and three atoms of hydrogen must be added to each of
the nitrogen atoms. The chemical process of nitrogen fixation involves "burning" of fossil
fuels to obtain the electrons, hydrogen atoms and energy needed to reduce molecular nitrogen.
Some bacteria are capable of N2 fixation from the atmospheric N pool. These bacteria
form various associations with plants:

(i) Many free living N-Fixing bacteria occur in the soil.

(ii) Some have adapted to form symbioses; others have intimate entophytic
associations with plants.

(iii) Others live in close association in the plant root zone (rizosphere) without
forming intimate endophytic symbioses (Rai, 2005).

Azotobacter, Beijerickia and Clostridium are considered as free living N-Fixing


bacteria while Azospirillum, Bacillus, Klebsiella and Pseudomonas are considered as
associated N-fixing bacteria (Triplett, 1996; James et al, 1997; Boddey, 2000; James, 2000).

Azotobacter is Gram negative bacteria, polymorphic i.e. they are of different sizes and
shapes. Their size ranges from 2-10x1-2.5 µm. Young cell possess peritrichous flegella and
are used as locomotive organs (Fig.4). They also produce polysacharides. It is sensitive to
acidic pH, high salts, and temperature above 35 oC. (www. 2).

Azospirillum is described also as Gram negative, rod-shaped, 1 µm in diameter, very


motile. Cells are about 1.0x3.5 mm in size single flagellum. Azospirillum utilizes glucose,
lactate, fructose, malate, pyruvate, fumarate as carbon source, reduce nitrate and does not
require biotin. (www. 3).

Fig. 4. Azotobacter chroococcum*

*(www. A)
Nitrogen fixing bacteria obtains the required energy from the oxidation of
carbohydrates which have been formed by the photosynthetic activity of green plants. The
energy for photosynthesis comes from sunlight.

The energy for bacteria is therefore indirectly derived from a universally available and
inexhaustible source. The direct source of energy (carbohydrate) for bacteria is therefore
potentially available wherever conditions permit the growth of autotrophic organisms.
However, the ability of a biological system to fix nitrogen is dependent on the presence of a
particular enzyme “nitrogenase”. This enzyme catalyzes the conversion of N2 into a reduced
form (ammonia combined with certain organic compounds) which can then be used for
growth by microorganisms and higher life forms.

Fig. 5. Azospirillum brasilense*

*(www. B)

The nitrogenase system consists of two different protein molecules (enzymes) which
must function together in the nitrogen fixing process. The ammonia resulting from fixation is
rapidly incorporated into certain amino acids, such as glutamine or alanine. The nitrogen may
then be transferred to other amino acids and nitrogen-containing compounds by a variety of
commonly occurring amino transfer reactions (Hubbell and Gerald, 2003).

Azospirillum and Azotobacter are plant growth promoting rhizobacteria (PGPR)


capable of affecting the growth and yield of numerous plant species, many of agronomic and
ecological significance (Bashan and Holguin, 1997; Bashan et al, 2004; del Amor et al, 2008).
These PGPR have no preference for crop plants or weeds, or for annual or perennial plants,
and can be successfully applied to plants that have no previous history of PGPR in their roots
(Dobbelaere et al, 2003).
Plant growth promoting rhizobacteria not only have been reported to enhance plant
growth directly by a fixation of atmospheric nitrogen that is transferred to the plant, but also it
have the ability to produce siderophores that chelate iron and make it available to the plant
root, solubilization of minerals such as phosphorus, and synthesis of phytohormones (Glick,
1995; Dobbelaere et al, 2001). Direct enhancement of mineral uptake due to increases in
specific ion fluxes at the root surface in the presence of PGPR has also been reported
(Bertrand et al, 2000).

Azospirillum sp. are known mainly for their ability to produce plant hormones as well
as polyamines and amino acids in culture (Thuler et al, 2003), but they are also involved in
the biological fixation of nitrogen and the increased activity of glutamate dehydrogenase and
glutamine synthetase (Ribaudo et al, 2001).

Azospirillum brasilense produces high quantities of extracellular indole acetic acid


which lead to increase root elongation, root surface area, and root dry matter (Molla et al,
2001). The effects of these microorganisms are influenced greatly by the species and
genotype (Sensoy et al, 2007), cultural practices such as fertilizer application (Gryndler et al,
2001). Although different studies have correlated plant growth and yield response with PGPR
inoculation, so far little is known about its specific effect on fruit duality

The amount of N fixed by these different microorganisms is considerable, although


variations resulting from environmental conditions or different plant microbe combinations
are vast. The close proximity of these microorganisms to their host plants allows efficient
plant use of fixed N and minimizes volatilization, leaching, and denitrification (Rai, 2005).

Table Sea 35 6.
Total Type of fixation N2 fixed (10135
12
g per year)

Non-biological
Industrial 50
Combustion 20
Lightning 10
Total 80
Biological
Agricultural land 90
Forest and non-agricultural land 50
Estimates of the amount of nitrogen fixed on a global scale*
*(Bezdicek and Kennedy, 1998)
3.2. Arbuscular Mycorrhiza
Mycorrhiza is involved in the most durable, intimate and important symbiosis on the
earth (Allen, 1996; Hodge, 2000), and form mutualistic symbioses with a vast majority of
land plants (Smith and Read, 1997). Possibly more than 80 % of all terestric plants form
mycorhizal symbioses.

Mycorrhiza is traditionally classified into two types, ectotrophic and endotrophic.


This classification was based on the position of the fungal hyphae in relation to the root
tissues of the plant (Raj, 2005). Ectomycorrhizal hyphae ensheath the root cells, but do not
penetrate them, i.e. they remain extra cellular. While, endomycorrhizal hyphae are
intracellular as they actually penetrate the root cells of plants.

This classification is now regarded as too simplistic, and has been superseded by a
nomenclature identifying 7 mycorrhizal types. The term ectomycorrhiza remains, as the fungi
within this group are reasonably homogenous, but a sub-group, ectendomycorrhizas, is added.
Within the endotrophic group, 5 distinct types have been classified as follows:

(i) Arbuscular mycorrhiza (AM),

(ii) Orchidaceae, based mycorrhiza

(iii) Mycorrhiza in the Ericales family, sub-divided in to Arbutoid, Monotropoid and


Ericoid mycorrhiza.

Arbuscular mycorrhiza association is endotrophic, and has previously been referred to


as Vesicular Arbuscular Mycorrhiza (VAM). This name has since been dropped in favor of
AM, since not all of the fungi form vesicles (Deacon, 1997).

The association between plants and their root colonizing mycoorhiza fungi is a
functional symbiosis in which the mycorhizal fungus is obligatory or facultative dependent on
host photosynthesis and energy. The symbiosis develops in the roots where the fungus
colonizes the cortex and obtains carbon from the plant while facilitating the transfer of
mineral nutrients to the root cells (Harrison, 2005).

The most important benefit of mycorrhiza is the increase in nutrient absorption, such
as H2PO4-, Zn++, Cu++ and NH4+. These benefits lead to improve the nutritional status of the
plants, to widen their ability to adapt to different ecosystems, and enhance the tolerance to
stressful biological and non-biological factors (Siqueira and Saggin Junior, 1995).
Thus, resulting from the more nutrient uptake, plant growth is generally improved
when mycorrhizal fungi colonize the root system (Rai, 2005). Moreover, when there is a
water deficit, the direct transportation of water to the roots can be interrupted and, in this case,
the external hyphae also acquire an important role in the absorption of water (Allen and Allen,
1986).

Fig. 6. Arbuscle of Glomus mosseae with numerous fine branch hyphae*

*(www. C)

Fig. 7. Mechanism of establishment of Endo-mycrrohiza symbiosis*

*(www. D)
Besides the nutritional benefits, numerous non-nutritional benefits have been
attributed to the AM, including:

(i) improvement of plant water relationship

(ii) higher tolerance to drought

(iii) higher tolerance to pathogens, salinity and heavy metals

These non-nutritional benefits can be related to alternations in the metabolism and


physiology of mycorrhiza plants (Schwab et al, 1991). Mycorrhiza fungi can cause
physiological alterations, increasing the stomatal conductance, respiration, transpiration,
absorption of CO2, photosynthesis (Mathus and Vyas, 1995; Ruiz-Lozano et al, 1995;
Goicoecheta et al, 1997; Rilling et al, 2002).

In addition, mycorrhizal plants can enhance the accumulation of carbohydrates


(Thomson et al, 1990), amino acids (Johansen et al, 1996) and cause alternations in the
content of hormones (Goicoecheta et al, 1997), and increase enzymatic activity (Ruiz-Lozano
and Azcon, 1996), as well as secondary metabolites (Peipp et al, 1997).

Also, the yield increase can easily be demonstrated in greenhouse experiments


(Johnson et al, 1997). Moreover, arbuscular mycorrhiza have become a tool to improve
seedling production before transplanting and help them to improve their establishment and
adaptation to field condition after they are transplanted (Rai, 2005).

All organic growing media contain microorganisms (Carlile and Hammonds, 2008).
The species and number of these microorganisms depends on several factors including the
constituents of media, their associated moisture and nutrient content, pH, C:N ratio and types
of carbon. Raw peat has a low microbial content, linked to its low pH and high lignin content
(Kavanagh and Herlihy, 1975).

In connection, Cordiki et al (2004) tested several different commercial AM inocula


and found that some were more efficient than others at developing mycorrhiza when mixed in
a peat based growth substrate.

Development of microbial numbers and activity in growing media containing peat is


linked to limits substrate, and especially when peat is combined with other organic materials.
Microbial activity is far more pronounced in many other organic materials used in the
manufacture of growing media. This is particularly true of composted materials since the
composting process it self is due to the degradative action of microorganisms.
Many studies (Dickinson and Carlile, 1995 and 2004) have shown much higher levels
of microbial activity in growing media based on composted materials. This microbial activity
in some circumstances are beneficial (Raviv, 2008) or detrimental in the case of
microorganisms that may cause nitrogen immobilization and/or those that attack plants.

A shift of perspective occurred when Carlile and Wilson (1991) presented their review
on microbial activity in growing media which highlighted the influence of microorganisms
associated with growing media regarding their physical and chemical qualities. They
concluded that this manipulation of microbial growth in soilless culture may offer
considerable benefits to growers and may form an integral part of commercial use of soilless
media in a near future (Al-Moshileh, 2007).

Moreover, inert growth systems such as rockwool are deficient in microbial activity,
and applying mycorrhiza inocula to these systems may not only result in microbial
mycorrhiza formation, but also in introduction of mycorrhiza associated bacteria (Mansfeld-
Giese et al, 2002). In this respect, Bin et al (2008) found that mycorrhiza associated bacteria
from the genus Paenibacillus have been shown to promote growth of cucumber plants.

As mentioned before, the effects of different microorganisms inoculation will be


reviewed as follows in the next chapters.

3.3. Vegetative growth characteristics


Various bacteria can promote plant growth (Bashan, 1998). Many researchers found
that N-Fixers have positive effects on the vegetative growth characters of many vegetable
crops. In this regard, Shahaby and Agwa (1993) revealed that inoculation of spinach seeds
with Azotobacter significantly increased fresh and dry weight of plants.

In connection, Gomma (1995) reported that single inoculation with Azotobacter


significantly increased cucumber plant fresh weight by 17 %, while no significant differences
in plant length and dry weight were reported among the individual inoculation including
Azotobacter and the control treatment (Hoagland nitrogen solution).

In comparison among different N-Fixing bacteria, Gupta et al (1995) pointed out that
inoculation tomato plants with Azotobacter was most effective in increasing the total dry
weight, root and shoot length, followed by Azospirillum.
Indeed, several studies indicated that Azospirillum can increase the growth of various
crops. These include carrot, eggplant, pepper, oak, sugar beet, sugarcane and tomato (Bashan
et al, 1989; Bashan and Holguin, 1997).

In this respect, Barakat and Gabr (1998), under inoculation with Azospirillum,
recorded a significant increase in the tomato seedling height, leaf area, fresh and dry weight,
but not in number of leaves per seedlings. In accordance, Cho Ja Yong et al (1998) mentioned
that Azospirillum sp. promoted early growth of cucumber and tomato plug seedlings.

These results are in harmony with Tantawy (2000), who found that inoculation of
tomato seeds with Azospirillum increased stem length, plant dry weight and plant fresh
weight, which reached 13.5, 35 and 52 %, respectively. In connection, Garib (2002), found
that inoculation of cucumber with Azotobacter and/or Azospirillum promoted different growth
characters i.e. number of branches, number of leaves and leaf area under greenhouse
condition.

In addition, Polyanskaya et al (2002) mentioned that Beijerinckia mobilis and


Clostridium spp. stimulated growth of cucumber and barley plants under greenhouse
condition. Moreover, Nuruzzaman et al (2003) revealed that applications of Azotobacter or
Azospirillum alone or accompanied with 5t.ha-1 cow dung fertilizer to okra resulted significant
variations between treatment in respect of morphological characters, e.g. plant height and
number of leaves per plant, and physiological characters like leaf area index (LAI) and crop
growth rate. Also, Wahba et al (2004) found that Azospirillum lipoferum affected positively
growth of carrot and turnip crops.

These positive effects resulted from bacteria inoculations are related to their ability to
improve plant nutrition by fixing atmospheric N, and influence plant growth by production of
plant enzymes (Gruda et al, 2004), hormones such as gibberellins (Gutierez-Manero et al,
2001) and cytokinins (Timmusk et al, 1999).

In connection, Miles (2007) reported that Azotobacter is able to synthesize auxins,


vitamins, growth substances and antifungal antibiotics which lead to beneficial effects on seed
germination of several crops.

Similar suggestions were documented by Bashan and Holguin (1997) whom


mentioned that genus Azospirillum can fix atmospheric N2 and enhance plant mineral uptake
as well as produce plant growth promoting hormones.
However, Bohme et al and Juhlke et al (2005) reported that better plant growth due to
the addition of microorganisms could sometimes be obtained in the soilless culture and in
container plant production.

On the other hand, Gomma (1995) revealed no response of tomato seedlings, as plant
height, fresh and dry weight of plants, if inoculated by Azotobacter and Azospirillum. Similar
trend was mentioned by Gruda et al (2008), whom observed that additives of Bacillus subtilis
did not affect fresh and dry weight, number of leaves as well as leaf area of primula and
begonia plants.

However, it is difficult to explain the role of Bacillus subtilis since a little has been
published on the physiological mechanisms involved as opposed to the agronomic effect of
this microorganism (Saleh et al, 2005).

Regarding to the effect of arbuscular mycorrhizal (AM) fungi on plant growth,


Changxian Wang et al (2006), indicated that growth of cucumber seedlings was significantly
enhanced by Glomus mosseae, inhibited by Glomus versiforme, and did not significantly
influenced by Glomus intraradices, as well as dry weight of the inoculated seedlings was 20
% times higher than control.

Lu GuiYun et al (2006), reported that G. versiforme, G. mosseae, G. intraradices and


their mixtures, which applied to cucumber seedlings in greenhouse increased plants leaf area,
and enhanced the healthy seedling index (HSI) by 58 % - 124 % if compared to control (Lu
GuiYun et al, 2002).

On the other hand, Douds and Reider (2003) found that inoculation of sweet pepper
seedlings with G. intraradices prior to transplanting decreased relative seedling growth to
control. Sabin and Larsen (2005) did not reveal differences between mycorrhizal and non-
mycorrhizal tomato plants in terms of growth parameters.

Moreover, Yesilova and Karaca (2007) mentioned that inoculation of melon plants
with G. entunicatum increased significantly plant and root lengths as well as shoots dry
weight when compared with control plants under the laboratory condition, while plant growth
parameters were not significantly differ under greenhouse conditions. Recently, application of
mycorrhiza to pepper plant improved plant growth than control under soilless culture
condition (Ikiz et al, 2009).
3.4. Yield and its component
In two decades of field experiments, general consensus is that in 60 to 70 % of the
cases Azospirillum application results in a significant crop yield increase (Okon and
Labandrea-Gonzalez, 1994).

Yield increase can be substantial, up to 30 %, but it generally ranged from 5 to 30 %.


These yield increases by Azospirillum are possibly a result of the production of growth
promoting substances rather than N2 fixation (Okon, 1985). In connection, Shahaby et al
(1993) noticed that inoculation of tomato seedlings with biofertilizers containing Azotobacter,
Azospirillum and/or Clostridium significantly increased fruit yield.

In addition, Li Ning et al (1997) and Grelalorenzo et al (1988) found that applying N-


Fixers increased fruit set and yield of pumpkins and tomato, respectively. Therefore, Omar
and El-Kattan (2003) revealed that inoculation of cucumber cv. Passandra with Bacillus
polymyxa as N2-fixing bacteria increased significantly total yield, by 15 % over un-inoculated
control.

Meanwhile, Panwar et al (2000) mentioned that inoculation of Azotobacter +


Azospirillium + 120 kg N/ha recorded the highest seed yield and values quality for radish. In
accordance, Wahba et al (2004) cleared that Azospirillum lipoferum affected positively yield
of carrot and turnip.

In this respect, Miles (2007) mentioned that Azotobacter increase plant yield primarily
by fixing molecular nitrogen, but it is also reported to synthesize auxins, vitamins, growth
substances and antifungal antibiotics, which have beneficial effects of this bacterium on seed
germination. The author revealed that Azotobacter spp. inoculants can increase crop yields on
barely, cabbage, maize, oat, onion, potato, rice, sugarcane, tomato, wheat by 7-12 %.

Regarding to the effect of arbuscular mycorrhizal (AM) fungi inoculation, Douds and
Reider (2003) found that inoculation of sweet pepper seedlings with Glomus mosseae prior to
transplanting showed significant increase in fruit yield, while Glomus intraradices decreased
yield relative to control.

Moreover, Qi GuoHui et al (2001) found that Glomus versiforme, Glomus mosseae,


Glomus intraradices increased fruit yield, number of fruits, early and total yield compared to
control in strawberry. Similarly, weights of single cucumber fruit of plants inoculated with G.
mosseae and G. versiforme were about 1.4 and 1.3 times higher than those from the un-
inoculated treatment, respectively, (Changxian Wang et al, 2006). In sweet potato, Glomus
mosseae led to a significant increase in tubers yield when compared to not inoculated plants
(Novak et al, 2007).

In contrast, Tantawy (2000) observed that un-inoculated tomato plants (control),


which received NPK fertilizers, produced significantly the lowest early, marketable and total
yield of tomato. In addition, Gul et al (2007) did not find significant differences between
inoculated tomato plants with Bacillus subtilis and control plants in terms of total yield.

In this respect, Sabin and Larsen (2005) studied the possibility of implementing AM
fungi in commercials cucumber production. The authors found, in the first production cycle
from January to July in two greenhouses, that AM did not influence cucumber yield, but in
the third nursery inoculation with AM resulted in approximately 4 % more in first class
cucumber compared with plants without mycorrhiza, which would correspond to an extra
profit of approximately 50,000 Euro/ha in the spring production cycle.

3.5. Chemical component


According to the effect of microorganisms on fruit quality, Qi GuoHui et al (2001)
working on strawberry found that AM fungi Glomus intraradices, Glomus mosseae, Glomus
versiforme increased ascorbic acid and sugar contents, while the titratable acid content in the
fruits was reduced by inoculation treatments, and among the AM fungi, G. versiforme was the
most effective fungi.

Recently, Lu GuiYun et al (2006) reported that G. intraradices, G. mosseae, G.


versiforme and their mixtures, which applied to greenhouse cucumber seedlings, could
significantly improve the quality of cucumber fruits.

Moreover, Chaurasia et al (2001) declared that soil application of Azotobacter at 15


kg.ha-1 coupled with N, P and K at 150, 60 and 80 kg.ha-1 influenced positively total soluble
solid and shelf life of eight tomato hybrids, while phosphate solubllizing bacteria did not
influence significantly tomato quality.

On the other hand, Gul et al (2007) documented that inoculation of tomato with
Bacillus subtilis or Trichoderma horyianum did not affect vitamin C or fruit quality
characteristics.

Regarding to the effect of microorganisms on mineral composition, Shahaby and


Agwa (1993) revealed that inoculation seeds of spinach with Azotobacter chroococcum alone
or plus Azospirillum brasilense increased significantly total nitrogen, as well as the uptake of
phosphorus, iron and potassium in spinach leaves. Also, Poi (1998) found that inoculation
with Azospirillum and phosphate dissolving bacteria (PDB) stimulated a higher uptake of N
and P in tomato and okra.

In connection, Demir (2004) mentioned that leaves of pepper plants inoculated with
Glomus intraradices showed better phosphorus, dry matter, chlorophyll and amount of some
reducing sugars (fructose, alpha glucose and beta glucose) comparing to un-inoculated plants.
These results are in harmony with Neumann (2005) who reveled that AM fungal inoculation
increased phosphorus uptake of tomato cv. Golden Queen.

In cucumber, nitrogen, chlorophyll and dry matter content in leaves were increased
under greenhouse conditions using combination of Azospirillum + Azotobacter + phosphate
dissolving bacteria (Garib, 2002).

In addition, in two pot experiments, cucumber (Cucumis sativus L. cv. Jinlu No. 3)
seedlings were each inoculated with one of three arbuscular mycorrhizal fungi, Glomus
mosseae, Glomus intraradices and Glomus versiforme. Contents of nitrogen and phosphorus
in roots and magnesium, copper, and zinc in shoots were increased by inoculating treatments,
while potassium and iron contents in shoots decreased significantly (Changxian Wang et al,
2006).

III. MATERIALS AND METHODS


This experiment was carried out during the two successive seasons of 2006 and 2007
under the controlled greenhouse condition at Faculty of Horticulture in Lednice, Mendel
University, Czech Republic.
The objective of this work was to study the effect of inoculation with nitrogen fixing
bacteria (Aztobacter chroococcum and Azospirillum brasilense) as well as arbuscular
mycorrhiza fungi (Glomus mosseae) on vegetative growth, yield, fruit quality and chemical
composition of cucumber plants grown on rockwool and peatmoss, as based growing media,
in an open soilless system.
Seeds of cucumber (Cucumis sativus L. cultivars ‘Passandra’ and ‘Girola’) were sown in
trays filled with sphagnum peat on 23rd and 19th of January 2006 and 2007. One week later, in
the first proper leaf stage, seedlings with complete root cone were shifted to rockwool cubes
(75 mm) and moved to growing tables till transplanting.
Transplants, at 2-3 true leaves stage, settled in the greenhouse on 7th of March 2006 and
19th of February 2007, respectively.
Two plants were placed in one plastic bag (1000 x 120 x 70 mm) filled with rockwool or
5 kg of peatmoss as based growing media. Plant spacing was 0.5 x 0.5 m. The plastic bags
were placed in double rows and each plant had its own irrigation capillary.
Irrigation system assured by DGT Volmatic technology. The climatic data (temperature,
sun radiation, air humidity) were evaluated as a factor influencing plant growth.
Fertigation doses were applied according to the plants requirements and climate
conditions. The plants were fertigated with a nutrient solution of following composition
(mg.L-1): 5 NH4+, 185 NO3-, 45 P, 200 K, 275 Ca, 45 Mg, 0,6 Fe, 0.3 Zn, 0.3 Mn and 0.3 B.
The electrical conductivity of the nutrient solution was 2.5 mS.cm-1 and the pH was adjusted
to 6.0 using HNO3 and dilution or addition of nutrient solution. Hydroponic system was
characterized and evaluated by regular analysis of nutrient solution composition in root-zone
area (minerals content, pH, conductivity).
Greenhouse environment and crop management practices were in accordance with
standard recommendations for commercial growers.

Table. A. Mean mineral content of nutrient solution in growing media

2006 2007
Content
Peatmoss Rockwol Peatmoss Rockwol

EC (mS.m-1) 1,8 2,3 2,6 3,0

pH 5,5 6,3 5,9 6,5

Ca (mg.l-1) 310 196 340 280

K (mg.l-1) 130 144 240 432

Mg (mg.l-1) 75 19 35 57

NH4+ (mg.l-1) 2,9 3,2 3,0 3,9

NO3- (mg.l-1) 130 280 220 155

PO4- (mg.l-1) 20 28 9 13

This experiment included 32 treatments of combination of two cultivars grown on two


substrates, rockwool and coarse peat, in addition to seven microorganisms inoculation
treatments, as follows:
Cucumber F1’s cultivars

o Passandra F1
Introduced by Enza-Zaden company, Holland. It has good growing vigorous open plants,
giving side-shoots, 2-3 fruits/node and top yielding. The plants and fruits are bitter free. The
fruits are dark green, slightly ribbed and cylindrical with a length of 150 mm “mini fruit
type”. It is powdery mildew and scab tolerant as well as recommended for autumn and spring
crops.
o Girola F1
Introduced by Enza-Zaden company; Holland. It has strong vigorous plants, produces
high early and total yield. The fruits are dark green, smooth, slightly ribbed with a length of
330 mm “long fruit type”. It is leaf-spot, scab resistance and recommended for autumn as well
as winter planting.

Microorganisms inoculation
Nitrogen fixing bacteria Aztobacter chroococcum CCM 1921 and Azospirillum
brasilense CCM 3862 were obtained from Czech Collection of Microorganisms (CCM)
institute, Masaryk University, Faculty of Science, Brno.
Bacteria were applied to transplants, grown in 80 mm trays filled with sphagnum peat as
a transplants growing media, using liquid culture from each strain at rate of 1 ml liquid
culture per plant (1 ml contains 109 cells of bacteria) as described by Valentine et al (2002).
Arbuscular mycorrhiza fungi Glomus mosseae were mixed to sphagnum peat as a
growing media at rate of 3 g per plant as recommended by Symbio-m Company (CZ).
Microorganisms applications were replicated 2 times during transplants growing stage.
The first inoculation dose was applied around root zone at 14th of March 2006 and 26th of
February 2007, respectively. Second inoculation dose was applied 15 days after the first
treatment.
The inoculation treatments and its mixtures were applied as follows:
1. N1 (Azotobacter chroococcum)

2. N2 (Azospirillum brasilense)

3. G (Glomus mosseae)

4. N1 + N2 (A. chroococcum + A. brasilense)

5. G + N1 (G. mosseae + A. chroococcum)


6. G + N2 (G. mosseae + A. brasilense)

7. G + N1 + N2 (G. mosseae + A. chroococcum + A. brasilense).

Training system
The basic principles of training the plants to uniformities distribute the foliage
throughout the entire greenhouse so that the leaves can intercept maximum light.
Single crop wires was placed 2.5 m above the ground over each row of plants. The
strings for plants were then tied alternately to the overhead wires so that the plants were
inclined away from the row on each side. The plants let to grow in a V- arrangement down the
row, allowing fruit to hang away from the main stem.
Pruning
The pruning method has been prune to result in early and high fruit production as
described by Vogel et al (1996) as following:
1. Plants were trained to be overhead wire and the growing point was removed tell
500 mm for cultivar ‘Passandra’, as a mini fruit type, and 700 mm for cultivar
‘Girola’, as a long fruit type.
2. A small loop of string was tied around the wire and the main stem below the top
leaf to prevent the plant from slipping down the string.
3. All side shoots were cut after producing one fruits at a very early stage, except
one fruit - one leaf was left for cultivar Girola. While with cultivar Passandra, it was
allowed to produce one more fruit and one more lateral leaf after the first fruit has
been trained.
4. Three or four leaves have been selected when the main stem reached the top of
wire, then other 2 branches were allowed to grow down. Only the laterals were
topped when necessary (when they touch the ground).
Data recorded
I. Vegetative growth characteristics
Three vegetative growth plant samples were collected randomly from each replicate after

60 days after transplanting to determine the following growth characteristics:

1. Plant height.
2. Numbers of leaves/plant.
3. Total leaf area/plant.
4. Fresh and dry weights of leaves.
II. Yield

Harvest, at marketable green mature stage, started on 4th and 15th April and finished on 18th

and 5th June 2006 and 2007, respectively. The following yield parameters were recorded:

1. Early yield, the sum of the first five pickings (kg.m-2).


2. Total yield, the all harvesting pickings (kg.m-2).
3.
III. Fruit quality

To estimate fruit quality, samples of five fruits at the marketable green stage of the tenth

and ninths picking of the first and second season, respectively were chosen randomly from

each replicate, where the following properties were determined:

A. Physical properties

1- Fruit length and diameter were estimated by Vernier Caliper instrument in mm.
2- Average weight of fruit (g).
3- Quality according to the EU norm No. 46 3155.
B. Chemical properties

o Dry leaves and fruits:


A sample of 3 leaves and 5 green mature fruits from each replicate was taken after 60
days from transplanting. Leaf samples were taken from plants with fully expanded
leaves of similar size (fifth leaf). Fruits were removed from a zone at about two-thirds
of the total height of the plant. Leaves and fruits (not peeled) were rinsed three times in
distilled water after decontaminating with 1 % non-ionic detergent and then blotted on
filter paper. These samples were oven-dried for thee days and ground with motorized
miller.
1. Nitrogen was estimated using coulometry Eca Flow 130 GLP (ISTRAN
Com., SK). The fine powder (1.0 g) of dry sample was digested by using
sulphuric acid and perchloric acid according to Piper (1947). Dry samples
were mineralized before analysis using the micro-Kjeldahl method as
described by Jackson (1973).
2. Phosphorus was extracted in acidic medium, in the presence in vanadat and
molybdate will make a yellow color. Complex intensive color was measured
at 430 nm wave length, using spectrometer Jenway 6100 (GB) as described
by (Holm, 1954).
o Fresh leaves and fruits:
3. Potassium, magnesium and calcium using capillary isotachophoresis
IONOSEP 900.1 (Recman Com., CZ) (Boček et al, 1986).
o Fresh fruit tissues:
4. Vitamin C, using chromatography HPLC. (ECON methodic, ECON CZ)
5. Total soluble solids, using ABBE refractometer, AR2 (AKRUS Com., D).
6. Total sugars, as sum of D-glucose and D-fructose converted into D-glucose-
6-phosphate. In the presence of diaphoreses the NADH is formed to a blue
formazan form, which was determined reflectometrically using RQflex
reflectometer, (Merck Com., D).
7. Nitrate, using Ionic selective Electrode (ISE). (Lutounská and Pichl, 1983)
8. Content of dry matter at 100 °C.
In this experiment a split-split plot design in randomized complete blocks with four
replicates was adapted. The cultivars were assigned to the main plots while, substrates types
were occupied as subplots, as well as microorganisms inoculation treatments were arranged as
sub-subplot. All data were subjected to the statistical analysis and means were analyzed using
the analysis of variance (ANOVA). The differences between the means was determined using
the L.S.D. test at significant level p<0.05. Statistical package Unistat (Unistat, Inc., USA) was
used.
IV. RESULTS AND DISCUSSION
IV. 1. Vegetative growth characteristics
1.1. Effect of cultivar
Data in Table (7) reveal that cultivar Passandra recorded higher significant increase in
plant height (1.88, 1.42 m), number of leaves per plant (37, 28), leaf fresh (29, 24 g) and dry
weight (3.2, 2.3 g) of cucumber plants in 2006 and 2007 seasons, respectively, if compared to
cultivar Girola.
Moreover, no significant effects were observed in leaf area values between the two
cultivars in the first season, while cultivar Girola showed higher significant increase (1117
cm2) in the second season than cultivar Passandra (725 cm2). This could be related to the
difference in night temperature between the two seasons.
The present study revealed that plant growth was affected by cultivar type. It was clear
that Passandra cultivar, as mini fruit type, was exceeded Girola cultivar, as long fruit type, in
plant height, number of leaves as well as leaf fresh and dry weight. According to the produced
seed company, Enza-Zaden, cultivar Passandra is recommended as autumn and spring crop,
while cultivar Girola as a winter crop. It is well known that the vegetative growth of cultivars
resistant to low temperatures are strongly affected by higher temperatures in spring season.
This was previously proved in cucumber (Medany et al, 1999; Garib, 2002)

1.2. Effect of substrate


Soilless culture as a popular production system for vegetable crops contributes to
enhance growth, yield and fruit quality due to good control of the root environment (Van
Winden, 1988). In commercial production, type of substrates and nutrient applications are
very important factors affecting the production and quality of vegetables under open soilless
system.
Results in Table (7) show that cucumber plants grown in peatmoss presented higher
significant increase in number of leaves per plant (39, 27) and leaf dry weight (3.3, 2.2 g) in
both seasons, respectively, when compared to plants grown in rockwool.
These results are in harmony with Shahinrokhsar and Tavusi (2008) whom revealed
that number of leaves, fresh and dry weight of strawberry plants grown in coco peat were
significantly higher if compared to those grown in bark + perlite substrate.
Leaf area and leaf fresh weight showed different trends in both growing seasons. First
season showed significant increase in leaf area of cucumber plants grown in peatmoss (939
cm2) while no significant differences were detected in terms of leaf fresh weight.
Meanwhile, in the second season, cucumber plants grown in rookwool showed higher
significant increase of leaf area (1241 cm2), while plants grown in peat moss resulted
significant increase in leaf fresh weight (27 g).
These unstable trends were previously obtained by Inden and Torres (2004). The
authors declared that fresh and dry weights of tomato were significantly higher in plants
grown in perlite plus carbonized rice hull, while no differences in fresh and dry weights of the
4th leaf were recorded between plants grown in rockwool or coconut coir substrates.
From the above results, it can be observed that vegetative growth characteristics of
cucumber plants were positively affected by peatmoss, as based growing media, than
rockwool.
According to the effect of substrates, Fascella and Zizzo (2005) mentioned that
organic substrates have higher holding water and cation exchange capacity than rockwool.
Simultaneously, the lower aeration condition in rockwool, compared to coarse peat, causes a
decrease in O2 levels in rooting media which reflected on root activity, nutrient uptake
absorption and growth vigor (Holtman et al 2005).
In this respect, it can be suggested that these obtained differences in growth
characteristics of cucumber plants could be related to distinction in chemical and physical
properties of rockwool and peatmoss.
On the other hand, plant height did not record significant differences between the used
substrates (rockwool or peatmoss) in both growing seasons. Similar results were obtained by
Cantliffe et al (2003) who pointed out that type of growing media (perlite or pine bark) did
not affect stem length in pepper plants.
In connection, Abul-Soud et al (2003) found that plant height of cucumber plants
grown in pumice or perlite did not show any differences among the substrates. Meanwhile,
AL-Rawahy et al (2009) mentioned that plant height and stem thickness for cucumber plants
grown upon wood straw and date palm leaf straw in open soilless system did not show
differences between the two substrates.
In this respect, it can be suggested that non-significant differences might be not related
to the type of used substrates, but might be due to other factors, i.e. genotype. (Hanafy Ahmed
et al 2004; Abdelaziz et al, 2007).
1.3. Effect of microorganisms inoculation
Table (8) clears that inoculation of cucumber plants by N-Fixing bacteria and/or
Glomus did not affect vegetative growth characteristic in both growing seasons if compared to
control treatment, with some exceptions.

Table 8. Effect of microorganisms inoculation on some vegetative growth


characteristics of cucumbers in 2006 and 2007

Plant height Leaf area No. Leaves/ Leaf FW Leaf DW


Treatments
m cm2 plant g/leaf g/leaf
Season 2006
N1 17,5 904 36 26 2,9
N2 17,6 895 35 29 3,3
N1+N2 18,6 893 34 22 2,6
G 17,6 930 35 25 2,9
G+N1 18,0 877 37 29 3,4
G+N2 18,0 899 40 25 2,8
G+N1+N2 17,8 910 33 26 3,2
Control 17,9 933 36 26 2,9
L.S.D. NS NS 2,3 2,8 0,4
Season 2007
N1 13,4 997 26 25 2,1
N2 12,9 843 26 24 2,1
N1+N2 13,2 1056 26 22 1,9
G 13,3 919 27 21 1,8
G+N1 13,7 944 27 23 2,0
G+N2 14,0 1033 27 23 1,9
G+N1+N2 13,3 789 26 23 1,9
Control 13,3 784 27 22 1,8
L.S.D. NS NS NS NS NS
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
In the first season, inoculation of cucumber plants by Glomus + Azospirrillum
recorded significant increase in number of leaves per plant (40) if compared to control (36), as
shown in Fig. (8). Also in Fig. (8), inoculation of cucumber plants with Azospirrillum or
Glomus + Azotobacter resulted similar significant increase in leaf fresh weight (29 g) than
control (26 g).
The same combination (Glomus + Azotobater) led to a significant increase in dry leaf
weight (3.4 g) than control (2.9 g), as shown in Fig. (9).

Fig. 8. Effect of microorganisms inoculation on number of leaves and


leaf FW of cucumbers in 2006

50

40

30

20

10
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

No. leaves per plant Leaf FW (g/leaf)

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Many investigators found that inoculation of vegetables by different microorganisms


did not affect growth characteristics. In this respect, Gomma (1995) revealed no response of
tomato seedlings, as plant height, fresh and dry weight of plants, if inoculated by Azotobacter
and Azospirillum. In connection, Barakat and Gabr (1998), under inoculation with
Azospirillum, recorded no significant increase in number of leaves of tomato seedlings.
Recently, Gruda et al (2008) observed that application of Bacillus subtilis did not
affect fresh and dry weight, number of leaves as well as leaf area of primula and begonia
plants.
Fig. 9. Effect of microorganisms inoculation on leaf DW of cucumbers
in 2006

4,0

3,5
g/leaf

3,0

2,5

2,0

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Concerning role of microorganisms on plant growth, Hubbell and Gerald (2003)


mentioned that nitrogen fixing bacteria obtains the required energy from the oxidation of
carbohydrates which have been formed by the photosynthetic activity of green plants.
However, it is well known that energy for photosynthesis comes from sunlight.
In this respect, it can be suggested that limited light quantities inside greenhouse in
winter season might cause a decrease in carbohydrates production. Thus, decrease the
function of nitrogen fixing bacteria. In addition, the variation in pH and saltation of the
nutrient solution could inhibit the microorganisms formation under such conditions (Hanafy
Ahmed et al, 2002 c).
On contrast, many researchers found that N-Fixers have positive effects on the
vegetative growth characters of many vegetable crops (Tantawy, 2000 on tomato; Garib, 2002
and Polyanskaya et al, 2002 on cucumber and Nuruzzaman et al, 2003 on okra as well as
Wahba et al, 2004 on carrot and turnip).
These positive effects based on fact that Azospirillum and Azotobacter, as plant
growth promoting bacteria, are able to produce plant hormones, polyamines and amino acids
in growing culture (Thuler et al, 2003). In connection, Gruda et al (2004) mentioned that
production of enzymes by microorganisms could be the reason for better growth.
In addition, the ability of those bacteria to fix small amount of nitrogen led to
encourage cell division and cell elongation (Meyer and Anderson, 1952). These positive
effects of N-Fixers reflected in an increase in plant height, number of leaves per plant as well
as fresh and dry weights of sweet pepper leaves (Hanafy Ahmed et al, 2002c).
Regarding to the effect of arbuscular mycorrhizal (AM) fungi on plant growth, Sabin
and Larsen (2005) did not detect differences between mycorrhizal and non-mycorrhizal
tomato plants in terms of growth parameters.
Meanwhile, Changxian Wang et al (2006), indicated that growth of cucumber
seedlings was significantly enhanced by Glomus mosseae, inhibited by Glomus versiforme,
and did not significantly influenced by Glomus intraradices.
In addition, Yesilova and Karaca (2007) revealed that inoculation of melon plants with
G. entunicatum increased significantly plant and root lengths as well as shoots dry weight
when compared with control plants. Recently, application of mycorrhiza to pepper plant
improved plant growth than control under soilless culture condition (Ikiz et al, 2009).

1.4. Effect of interactions


As shown in Table (7), the interaction between cultivar x substrate in the second season
reveal significant increase in plant height of cultivar Passandra (1.43 cm) and leaf area of
cultivar Girola (1588 cm2) grown in rockwool. Moreover, both cucumber cultivars recorded
the highest significant increase in leaf fresh weight (25 g) when grown in peatmoss than
control, in the same season.
In this respect it can be concluded that interaction between cultivars x substrate did show
preferable effect on most of growth characteristics of cucumber plants. Similar results were
obtained by Cantliffe et al (2003) on papper and AL-Rawahy et al (2009) on cucumber.
According to the interaction between cultivar x microorganisms inoculation, data in
tables (9 and 11) did not show significant differences in terms of plant height and leaf area in
both seasons (Sabin and Larsen, 2005).
Moreover, number of leaves per plant recorded the highest significant increase (44)
with cultivar Passandra inoculated by Glomus + Azotobacter in the first season only (Table
13). This result is in harmony with Barakat and Gabr (1998) on tomato.
In addition, leaf fresh and dry weight of both cultivars positively responded to the
different inoculation treatments but without clear trend in the two seasons as shown in table
(15 and 17).
Regarding to the interaction between substrate x microorganisms inoculation, Tables
(10, 12 and 14) reveal that plant height, leaf area and number of leaves of cucumber plants
grown in peatmoss responded positively to microorganisms inocultion in the first season only,
if compared to plants grown in rockwool (Figs. 10-12).

Fig. 10. Effect of interaction between substrate x microorganisms inoculation


on plant height of cucumbers in 2006

2,2

2,0

1,8
m

1,6

1,4

l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Rockwool Peatmoss

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Similarly, plants grown in peatmoss and inoculated with Azospirillum showed higher
value of leaf fresh weight (31 g), when compared to other treatments in the first season (Fig.
13). No differences were found in the second season (Table 16).
While in Table (18), cucumber plants grown in peatmoss and inoculated with (Glomus
mosseae + Azotobacter) led to a significant increase of leaf dry weight (4 g) in the first
season, as shown in Fig. (14).
Fig. 11. Effect of interaction between substrate x microorganisms inoculation
on leaf area of cucumbers in 2006

1200

900
2
cm

600

300

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Rockwool Peatmoss

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig. 12. Effect of interaction between substrate x microorganisms inoculation


on number of leaves per plant of cucumbers in 2006

50

40

30

20

10
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Rockwool Peatmoss

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


Fig. 13. Effect of interaction between substrate x microorganisms inoculation
on leaf FW of cucumbers in 2006

40

30
g/leaf

20

10

l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Rockwool Peatmoss

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig. 14. Effect of interaction between substrate x microorganisms inoculation


on leaf DW of cucumbers in 2006

5,0

4,0

3,0
g/leaf

2,0

1,0
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Rockwool Peatmoss

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


In this respect, many researchers found that N-Fixers have positive effects on the
vegetative growth characters of many vegetable crops as plant growth promoting bacteria
(Bashan et al, 2004; del Amor et al, 2008). Enhancement of plant growth may occur by direct
fixation of atmospheric nitrogen which transferred to the plants, and synthesis of
phytohormons (Dobbelaere et al, 2001), as well as increase the specific ion fluxes at the root
surface (Bertrand et al, 2000). These increases are quiet expected since it is well established
that decreasing the C/N ratio within the plant usually favors vegetative growth (Hanafy
Ahmed et al, 2004).
In addition, Siqueira and Saggin (1995) found that Glomus mosseae increase micro
nutrients absorption of Zn++ and Cu++ as well as improve the nutritional status of the plants.
Thus, resulting from the more nutrient uptake, plant growth is generally improved when
mycorrhizal fungi colonize the root system (Rai, 2005).
Therefore, it can be concluded that inoculation with N-Fixers alone/or combined with
Glomus mosseae resulted better growth characteristics of cucumber plants if compared to
other inoculation treatments, especially when plants grown under peatmoss.
IV. 2. Yield and its component
Data concerning the influence of different cultivars, substrates and microorganisms
inoculation and their interactions on cucumber yield and its components, i.e. early, total yield,
fruit length, diameter and numbers of fruits per plant as well as average fruit weight are
presented in Tables (19-44).

IV. 2.1. Early and total yield


2.1.1. Effect of cultivar
Data in Table (19) show that early and total yield of cultivar Passandra (0.7 and 17.0
kg.m-2) presented light higher significant increase if compared to cultivar Girola (0.6, 15.8
kg.m-2) in the first season, respectively.
While, early yield of cultivar Girola (3.2 kg.m-2) were significantly higher than
cultivar Passandra (2.1 kg.m-2) in the second season.
In general, it can be observed that differences between early and total yield of both
cultivar are similar. In this respect, it can be suggested that type of cultivar did not affect early
or total yield of cucumber plants and cultivars are varied in their yield productivity (Arold,
1998; Broeck 1999; Graib, 2002)

2.1.2. Effect of substrate


Results in Table (19) reveal light significant increase in early yield of cucumber plants
grown under rockwool than peatmoss in the two seasons. On contrast, total yield of
cucumbers grown under peatmoss recorded higher significant increase (17.0 kg.m-2) if
compared to cucumbers grown in rockwool (15.8 kg.m-2), in the first growing season.
However, an exceeded variation in total yield between cucumbers was recorded in the second
season, as shown in Fig. (15).
Plants showed the best significant increase (13.0 kg.m-2) when grown under rockwool
comparing to plants grown under peatmoss (9.9 kg.m-2). These results might be explained
based on fact that rockwool able to absorb more nutrients than peat slabs.
Similar explanation was mentioned by Chohura and Komosa (2000), Benko et al
(2008) on tomato and Bohme et al (2008) on cucumber.
Moreover, many authors found that type of growing media (organic or mineral) did
not affect yield productivity in cucumber (Cantliffe et al, 2003; Schroeder and Sell, 2009),
cherry tomato (Holtmann and Kobryn, 2003; Inden and Torres, 2004) and watermelon
(Lopez-Galarza et al, 2005).
Fig. 15. Effect of substrate on total yield of cucumbers in 2006-2007

20

16
-2
Kg.m

12

8
2006 2007

Rockwool Peatmoss

2.1.3.Effect of microorganisms inoculation


According to the effect of microorganisms inoculation on yield of cucumber plants,
results in Table (20) did not show any significant difference in early or total yield between
inoculated and control plants in both growing seasons, except total yield in 2007.
Mixture of Azotobacter + Azospirillum showed the highest significant increase of total
yield (12.6 kg.m-2) in the second season if compared to the control plants (10.4 kg.m-2).
Similar results were found by Ouda (2000) and Tantawy (2000) on tomato as well as Garib
(2002) on cucumber.
However, all inoculation treatment revealed better, but not significant, results in terms
of total yield if compared to control treatment in both growing seasons. These improve in total
yield reached 8 and 21 % in first and second season, respectively.
The obtained trend were confirmed by Omar and El-Kattan (2003) whom revealed that
inoculation of cucumber cultivar Passandra with Bacillus polymyxa, as N-Fixing bacteria,
increased significantly total yield by 15 % over un-inoculated treatment (control), and 10 % in
tomato when inoculated with Azotobacter (Miles, 2007).
In this respect it can be suggested that these favorable effects of N-Fixers on yield of
cucumber plants could be related to their ability to produce growth promoting substances
(Okon, 1985), synthesize auxins, indol acetic acids and gibberelins (Garib, 2002), vitamins
Table 20. Effect of microorganisms inoculation on early and total
yield (kg.m-2) of cucumbers in 2006 and 2007

Early yield Total yield Early yield Total yield


Treatments

Season 2006 Season 2007

N1 0,50 16,8 2,67 11,4

N2 0,67 16,8 2,71 11,0

N1+N2 0,69 16,4 2,65 12,6

G 0,73 16,6 2,71 12,1

G+N1 0,63 16,8 2,58 11,4

G+N2 0,63 16,5 2,60 11,9

G+N1+N2 0,56 15,6 2,79 10,9

Control 0,54 15,5 2,70 10,4

L.S.D. NS NS NS 1,4
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae

and antifungal antibiotics (Miles, 2007). Furthermore, presence of plant growth promoting
rhizobacteria (PGPR) might not result in signs of visible plant growth but contribute to plant
yield by reducing or suppressing disease incidents. It may be more practical to say that PGPR
help plant either directly via visible growth promotion or indirectly via disease control (Rai,
2005).
In this respect, it can be suggested that microorganisms inoculation did not play a vital
role to suppress disease function in the first season. However, it can be said that dual
inoculation of N-Fixers was partly more active in control disease and improve yield
productivity in the second season.
2.1.4. Effect of interactions
Inoculcated plants of cultivar Passandra with Glomus recorded the highest significant
values of early yield (0.9 kg.m-2) if compared to other treatments in the first season only as
shown in Table (21). Other interaction treatments did not show significant differences in both
seasons, Tables (21-22).
In terms of total yield, cultivar Passandra grown in peatmoss revealed higher
significant increase (17.5 kg.m-2) in the first season, and 13.6 kg.m-2 in the second season
when grown in rockwool, as shown in Table (19).
Moreover, total yield of cultivar Passandra presented better significant increase when
inoculated with Glomus (18.7 kg.m-2) followed by those inoculated with Azotobacter (18.2
kg.m-2) in the first season (Fig. 16).

Fig. 16. Effect of interaction between cultivar Passandra x microorganisms


inoculation on total yield of cucumbers in 2006

20,0

17,5

15,0
-2
Kg.m

12,5

10,0
l
1

2
1

G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

According to the interaction between substrate x microorganisms inoculation, data of


total yield in Fig. (17) show that cucumber plants grown in peatmoss and inoculated with
Glomus + Azotobacter or Azospirillium, were significantly higher (18.5 kg.m-2) in the first
season than control ( 16.0 kg.m-2).
Fig. 17. Effect of interaction between peatmoss x microorganisms
inoculation on total yield of cucumbers in 2006

20,0

17,5
-2
Kg.m

15,0

12,5

10,0

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

This enhancement of yield could be attributed to increase of available nitrogen and


phosphorus us and effects of such nutrients on flowering and fruit setting. Similar suggestion
was obtained by Saito et al (1963) and Fisher (1969). They reported that earlier buds were
obtained with the application of high level of nitrogen. Grelalorenzo et al. (1988) found that
increasing level of applied nitrogen increased the number of flowers and fruit set of
commercial tomato cultivars. In this respect it can be concluded that cultivar Passandra was
more responded to produce higher early and total yield than cultivar Girola when grown under
the inoculated peatmoss substrate if compared to un-inoculated control.
This could be related to the vigor of cultivar Passandra and its ability to absorb more
nutrients from root zone. In connection, Hanafy Ahmed et al (2004) suggested that increase
sweet pepper fruit yield under sufficient supply of N and P might be attributed to sufficient
supply of assimilates from the leaves to the fruits as a result of, a) an increase in available leaf
assimilate supply to the fruits, b) an increase in potential sink strength of fruit, and / or c) an
increase in translocation capacity.
IV. 2.2. Yield component
2.2.1. Effect of cultivar
Table (25) shows that fruits of cultivar Girola presented significant increase in terms
of fruit length, diameter and weight if compared to fruits of cultivar Passandra in the two
growing seasons. On contrast, plants of cultivar Passandra produced higher significant
number of fruits per plant than cultivar Girola.
According to the producer, Enza-Zaden company, cultivar Girola is classified as long
fruit type with 280 mm length and 30-50 mm diameter, while cultivar Passandra as mini fruit
type with 150-180 mm length and 30-40 mm diameter. So, it is expected to find out
significant differences between both cultivars in such above components. In this respect, these
results could be explained regarding to the difference in geno-type of each cultivar.

2.2.2. Effect of substrate


Data in Table (25) declare that fruits of cucumber plants grown in rockwool revealed
lightly increase in fruit length and weight than those plants grown in peatmoss in both
seasons.
These results could be explained according to Wallach et al (1992), Da Silva et al.
(1993) and Naasz et al (2005). The authors described that hydraulic conductivity of peat
radically decreases on restricted ranges of water potential that could rapidly lead to very weak
water flows in the substrate and that are potentially limiting for the root environment if
compared to mineral substrate, i.e. rockwool.
However, no differences were observed in fruit diameter for cucumbers grown under
the two compared substrates. In accordance, Parks et al, (2009) found that no significant
effect of coir, perlite and rockwool on cucumber fruit diameter.
Meanwhile, cucumber plants grown in peatmoss produces more fruits in the first
season (Broeck 1999 and Peyvast et al, 2008), while plants grown in rockwool produced more
fruits in the second season (Table 25).
This obtained low yield of cucumbers grown in peat moss in the second season could
be explained according to the negative effect of low temperature on water prosperities of peat
moss. It’s well known that peat moss has more satisfactory aeration and water holding
capacity than rockwool (Cantliffe et al, 2003; Iapichino et al, 2007). Low temperatures cause
a decrease in plant transpiration, vital process and nutrients uptake.
In addition, low temperature decrease evatranspiration and peat moss shrinkage. This
negative effects influence water and air aeration which lead to a decrease in aeration that
could cut off the supply of oxygen to the root during growth (Lemaire et al, 2003). Moreover,
low oxygen aeration cause problems in flower production and/or increase flower drops under
such conditions.

2.2.3. Effect of microorganisms inoculation


Inoculation of cucumber plants with microorganisms did not significantly affect fruit
length, diameter and numbers per plant in both growing seasons (Table 26).

Table 26. Effect of microorganisms inoculation on physical fruit

characteristics of cucumbers in 2006 and 2007

Fruit length Fruit diameter Fruit Fruit weight


Treatments
(mm) (mm) number/plant (g)
Season 2006
N1 226 37,0 29,7 253
N2 227 36,2 30,3 242
N1+N2 225 37,6 28,6 248
G 229 36,6 30,0 245
G+N1 228 37,2 29,8 242
G+N2 228 36,6 30,1 237
G+N1+N2 221 37,6 29,5 235
Control 227 36,5 27,5 241
L.S.D. NS NS NS 11,0
Season 2007
N1 240 39,4 22,0 235
N2 239 42,4 21,0 230
N1+N2 227 41,0 23,3 242
G 230 40,8 23,0 240
G+N1 234 4,1 24,5 223
G+N2 235 4,1 23,0 236
G+N1+N2 220 4,0 22,5 224
Control 232 4,0 22,1 227
L.S.D. NS NS NS NS
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
On the other hand, data in Fig (18) presented significant increase in average fruit
weight when cucumber plants inoculated with Azotobacter (253 g) alone or when combined
with Azospirillum (245 g) if compared to control (241 g) in the first season only.
Meanwhile, in the second season, fruit weight showed higher value (242 and 240 g)
when inoculated with (Azotobacter + Azospirillum) or Glomus, respectively, if compared to
the control (227 g).

Fig.18. Effect of microorganisms inoculation on cucumbers fruit weight


in 2006

260

250

240
g

230

220
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

However, these values did not reach to significant level but pointed out that
microorganisms inoculation had positive effects on increase cucumbers fruit weight.
Same results were recorded by Shahaby et al (1993) and Miles (2007) on tomato, Li
Ning et al (1997) on pumpkin, Omar and El-Kattan (2003) on cucumber whom found that
inoculation vegetables crops with microorganisms enhanced yield and its component.
In this respect, it can be suggested that preplanting microorganisms inoculation
treatments had favorable influence on average fruit weight. This might be a result of N-Fixers
effects on release more available nitrogen in root zone.
In accordance, improved levels of nitrogen and other nutrients led to positive effects
on many vital processes, i.e. chlorophyll, enzymes, photosynthesis and endogenous hormones
synthesis which consequently affect growth and yield (Marschner, 1995, Hanafy Ahmed,
1997 and Garib 2002).
In general, combination of N-Fixers or the single inoculation of Glomus enhanced the
vital process of plant organs and caused that increase in average fruit weight of cucumbers.
However, these effects of microorganisms reflected positively on average fruit weight than
fruit length, diameter or number. In particular, physical component such as fruit length and
diameter are more correlated to geno-type and less responding to microorganisms inoculation.

2.2.4. Effect of interactions


Data in Table (25) show that interaction between cultivar x substrate did not affect
yield components in both seasons. The only exception was recorded with cultivar Passandra
which produced higher significant number of fruits when grown in rockwool (35) than
peatmoss (27) in the second season only, as shown in Fig (19).
Fig.19. Effect of interaction between cultivar x substrate on number of fruits
per plant in 2007

40

30

20

10
Passandra Girola

Rockwool Peatmoss

Meanwhile, both cucumber cultivars produced higher values of fruit weight when
grown under rockwool than peatmoss in the second season (Fig. 20).
Moreover, the interaction between cultivar x microorganisms inoculation or between
substrates x microorganisms inoculation did not affect fruit length, diameter and weight in
both growing seasons (Tables 27 to 30). The only exception found in fruit diameter of cultivar
Girola which recorded the highest significant increase (40 mm) with Azotobacter+
Azospirillum + Glomus inoculation if compared to the inoculation treatments (Tables 33).
Fig.20. Effect of interaction between cultivar x substrate on
cucumbers fruit weight in 2007

400

300

200
g

100

0
Passandra Girola

Rockwool Peatmoss

In terms of number of fruits per plant (Table 31), it can be observed that cultivar
Girola revealed higher significant increase (22) when inoculated with Glomus + Azotobacter
+ Azospirrillum than control (20.6). While, cultivar Passandra trended similarly and produced
significant increase of number of fruits per plant when inoculated with Glomus (41), if
compared to other treatments in the first season, as shown in Fig. (21).

Fig.21. Effect of interaction between cultivar x microorganisms inoculation on


fruit number per plant of cucumbers in 2006

50

40

30

20 c

10
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Passandra Giola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


Regarding to the effect of interaction between substrate x microorganisms inoculation
on number of fruits per plant, Table (32) records that cucumbers grown in peatmoss showed
higher significant increase (35) when inoculated with Glomus + Azospirillum followed by
Glomus + Azotobacter (33) comparing to other treatments (Fig. 22).

Fig.22. Effect of interaction between peatmoss x microorganisms inoculation


on fruit number of cucumbers in 2006

40

35

30

25

20
N1 N2 N1+N2 G G+N1 G+N2 G+N1+N2 Control

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


IV. 2.3. Marketable yield
According to the EU norm No. 46 3155, cucumber fruits of each cultivar physically
classified to four grades as following:
1. Extra
2. Class A
3Class B
4. Unmarketable fruits

2.3.1. Effect of cultivar


Data in Table (35) show that fruits of cultivar Passandra recorded significant increase
(9.9 kg.m-2) of grade “Extra” if compared to fruits of cultivar Girola (5.1 kg.m-2) in the second
seasons only. Furthermore, percentages of classified grades for both cultivars are presented in
Fig (A). On contrast, no differences were found between both cultivars in terms of class A, B
or the unmarketable fruits except cultivar Passandra which revealed significant increase of
unmarketable fruits (0.20 kg.m-2) in the second season if compared to fruits of cultivar Girola
(0.07 kg.m-2).
This might be attributed to the high produced fruits number of cultivar Passsandra, as
mini fruit type, than cultivar Girola, as long fruit type, which led to increase percentage of
unmarketable fruits.

Fig. A. Percentage (%) of different grades to total yield of fruits


in 2006 and 2007

100

75 Extra
Class A
50
Class B
25 Unmarketable

0
Passandra Girola 2006 Passandra Girola 2007
2006 2007
2.3.2. Effect of substrate
Fruits of cucumber plants grown in peatmoss showed higher significant increase (12.2
kg.m-2) of grade “Extra” in the first season if compared to fruits of other cucumber plants
grown in rockwool (10.2 kg.m-2), as shown in Table (35).
In connection, Parra et al (2009) found that marketable yield of tomato plants grown
in coir was significantly higher than those grown in rockwool under open system condition.
In this respect it could be suggested that high fruits number of plants grown in
peatmoss (Table 25) caused these significant increase if compared to plants grown in
rockwool. Similar suggestion was obtained by Colla et al (2003) on cucumber.
On contrast, in the second season, fruits of cucumbers grown in rockwool presented
higher significant increase (8.9 kg.m-2) of grade “Extra” comparing to cucumbers grown in
peatmoss (6.1 kg.m-2), as shown in Table (35).
However, other grades did not show differences among the two substrate in both
growing seasons, except fruits of cucumbers grown in rockwool which showed significant
increase of “class B” (0.6 kg.m-2), in the second season. However, Fig (B) presents the
percentages of classified grades for cucumber grown under both cultivars.
In this respect it can be suggested that type of substrate did not affect marketable yield
of cucumber plants grown under both substrates. These results are agreed with those obtained
by Holtmann and Kobryn (2003); Inden and Torres (2004); Lopez-Galarza et al (2005) on
tomato, Cantliffe et al (2003) on pepper and Show et al (2004) on cucumber.

Fig. B. Percentage (%) of different grades to total yield of substrates


in 2006 and 2007

100

75 Extra
Class A
50
Class B
25 Unmarketable

0
Rockwool Peatmoss Rockwool Peatmoss
2006 2006 2007 2007
2.3.3. Effect of microorganisms inoculation
Inoculating cucumber plants with different microorganisms improved the marketable
yield of inoculated plants than un-inoculated plants (control).
Mixture of N-Fixers (Azotobacter + Azospirillum) led to a significant increase in the
marketable yield of grade “Extra” in the first season (13.1 kg.m-2) followed by Glomus (12.1
kg.m-2) if compared to control (10.5 kg.m-2). Similar trend were obtained in the second season
although results did not present significance influence, as shown in Table (Table 36).

Table 36. Effect of microorganisms inoculation on marketable fruit


yield (kg.m-2) of cucumbers in 2006 and 2007

Treatments Extra Class A Class B Unmarketable

Season 2006
N1 11,4 4,23 1,04 0,17
N2 10,7 4,16 1,73 0,20
N1+N2 13,1 2,53 0,75 0,00
G 12,1 3,18 1,29 0,02
G+N1 11,8 3,67 1,09 0,23
G+N2 11,4 3,67 1,29 0,15
G+N1+N2 10,8 3,01 1,49 0,31
Control 10,5 3,76 1,19 0,02
L.S.D. 1,47 0,94 0,44 NS
Season 2007
N1 7,8 0,88 0,28 0,03
N2 6,9 0,81 0,44 0,20
N1+N2 8,1 1,34 0,25 0,27
G 8,1 0,64 0,51 0,15
G+N1 7,4 0,96 0,51 0,05
G+N2 8,1 0,83 0,36 0,06
G+N1+N2 6,9 0,89 0,59 0,11
Control 6,9 1,00 0,24 0,20
L.S.D. NS NS NS NS
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
This might be attributed to the beneficial effect of microorganism’s inoculation to
increase the uptake of available nutrient from N and P in root zone of cucumber plants, and
then express their effects on flowering and fruit setting.
In this respect, it can be concluded that marketable yield was positively affected by
microorganisms inoculation. Similar conclusion was obtained by Veeranna et al. (2000)
and Hanafy Ahmed (2004).

2.3.4. Effect of interactions


Cultivar Passandra grown in rockwool had significant increase in marketable fruit
yield (11.6 kg.m-2) if compared to other treatments in the second season, while no differences
were recorded in the first season, as shown in Fig. (23).
Moreover, no significant differences were found between interactions in terms of class
A, B and unmarketable fruits in both growing seasons, Table (35).

Fig.23. Effect of interaction between cultivar x substrate on marketable


fruit yield of cucumbers in 2007

16

12
-2
Kg.m

0
Passandra × Passandra × Girola × Girola × Peatmoss
Rockwool Peatmoss Rockwool

According to the interaction between cultivar x microorganisms inoculation, Table


(37) shows that fruits of cultivar Girola presented the best significant increase in marketable
fruit yield (13.5 kg.m-2) when inoculated with mixture of N-Fixers (Azotobacter +
Azospirillum) in the first season and when inoculated with Azotobacter (5.9 kg.m-2) in the
second season, (Fig. 24). It can be suggested that these results could be related to the low
production of other grades under the same condition which led to an increase in grade
"Extra".
Fig.24. Effect of interaction between cultivar Girola x microorganisms
inoculation on marketable fruit yield of cucumbers in 2006 and 2007

16

12
-2
Kg.m

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

These finding are confirmed in Tables (39-40). Data in those tables indicate that
inoculation of cucumbers with mixture of N-Fixers (Azotobacter + Azospirillum) produced
the lowest fruit yield of "Class A" in the first season. Moreover, no significant differences
were observed in terms of "Class B" or unmarketable fruits in both growing seasons (Tables
41-44).
In addition, higher significant increase in marketable fruit yield was found with
cultivar Passandra (Fig. 25) when inoculated with Glomus (14.0 kg.m-2) followed by
Azotobacter + Azospirillum (12.7 kg.m-2) in the first season and 11.5 kg.m-2 in the second
season. Meanwhile, in Table (38), cucumber plants grown in rockwool and inoculated with
mixture of N-Fixers (Azotobacter + Azospirillum) showed higher significant increase in
marketable fruit yield (13.1 and 9.7 kg.m-2) in both growing seasons, respectively (Fig. 26).
Moreover, cucumber plants grown in peatmoss and inoculated by Glomus +
Azotobacter (13.3 kg.m-2) or Glomus + Azospirillum (13.2 kg.m-2) revealed significant
increase in the first season if compared to control (11.2 kg.m-2), as shown in Fig. (27).
In the second season, Glomus inoculation treatment produced higher significant
increase of marketable fruit yield (7.2 kg.m-2) if compared to control (5.8 kg.m-2).
In general, it can be noticed that cucumber plants grown in rockwool found to produce
higher marketable fruit than those grown peatmoss, as shown in Table (38).
Fig.25. Effect of interaction between cultivar Passandra x microorganisms
inoculation on marketable fruit yield of cucumbers in 2006 and 2007

16

12
-2
Kg.m

4
N1 N1+N2 G G+N1 G+N2 G+N1+N2 Control

2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig.26. Effect of interaction between rockwool x microorganisms


inoculation on marketable fruit yield of cucumbers in 2006 and 2007

16

12
-2
Kg.m

4
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

These results may be interpreted according to Thomson et al (1990) whom mentioned


that mycorrhiza can enhance the accumulation of carbohydrates, amino acids (Johansen et al,
1996) and cause alternations in the content of hormones (Goicoecheta et al, 1997), and
increase enzymatic activity (Ruiz-Lozano and Azcon, 1996), as well as secondary metabolites
(Peipp et al, 1997) which reflected on marketable yield productivity.
Fig.27. Effect of interaction between between peatmoss x microorganisms
inoculation on marketable fruit yield of cucumbers in 2006 and 2007

16

12
-2
Kg.m

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


IV. 3. Mineral components
Nitrogen, phosphorus, potassium, calcium and magnesium consider the most
important nutrients affect plant growth and yield of vast plants. In this respect, the effect of
cucumber cultivars, substrates, microorganisms inoculation and their interactions on contents
of above nutrients in cucumber leaves and fruits is presented in Tables (45-56) and Tables
(57-68), respectively.

IV. 3.1. Mineral composition of leaves


3.1.1. Effect of cultivar
As shown in Table (45), contents of nitrogen, phosphorus and magnesium did not
show significant differences in leaves of cultivar Girola or cultivar Passandra in the two
growing seasons. In this respect, it can be concluded that type of cultivar has no effect on
accumulation of such nutrients in both growing seasons.
Potassium was significantly higher in leaves of cultivar Girola (1074 mg.kg-1) if
compared to cultivar Passandra (1010 mg.kg-1) in the first season.
On the contrary, leaves of cultivar Passandra presented higher significant content of
potassium and calcium (1917 and 243 mg.kg-1) than cultivar Girola (1729 and 227 mg.kg-1) in
the second season, respectively.
In this respect it can be suggested that the differences in EC of nutrient solutions
among the two growing seasons caused these differences in K and Ca contents in cucumber
leaves, as shown in Table (A), as shown in materials and methods chapter. In connection,
Sonneveld and Welles (2005) found that K in cucumber tissues was increased and the Ca
concentration was decreased by an increasing EC in the external solution.

3.1.2. Effect of substrate


Nitrogen, calcium and magnesium reveal different trends in both growing seasons, as
shown in Table (45). Nitrogen (4.45 g/100g dry matter), calcium (193 mg.kg-1) and
magnesium (136 mg.kg-1) increased significantly in leaves of cucumbers when grown under
rockwool in the first season. This might be attributed to high content of NH4+ form in
rockwool than peatmoss. In connection, Dubsky and Sramek (2008) reported that rockwool
contained high potential of nitrogen fixation (40 mg.L-1) if compared to peatmoss (-83 mg.L-
1
).
On contrast, in the second season, the nitrogen (3.71 g/100g dry matter), calcium (278
mg.kg-1) and magnesium (144 mg.kg-1) were significantly higher when grown under
peatmoss.
The above findings may be explained by the different in chemical and physical
properties of substrates and their interaction with nutrient solution composition and plant
nutrient uptake. In this respect, it can be assumed that low EC of peatmoss in the second
season (2.6 mS.cm-1) if compared to rockwool (3.3 mS.cm-1) caused this decrease in Ca and
Mg contents. Similar results were obtained by Sonneveld and Welles (2005).
According to the effect of substrate on contents of phosphorus and potassium in leaves
of cucumber plants, Table (45) declares that phosphorus showed higher significant content
when cucumber plants grown in peatmoss in the first season only, while potassium showed
similar trend in the second season. This decrease in phosphorus, in the second season, could
be caused by the higher pH of rockwool if compared to peatmoss as shown in Table (A). In
connection, Colla et al (2003) found that magnesium and phosphorus us concentration was
higher in cucumber plants grown in coir than rockwool.
However, type of substrate affected nutrients content in the growing media, hence
accumulated in leaves of grown plants. Therefore, it can be concluded that optimal pH and
EC of rockwool than peatmoss led to release more quantities of N, Ca and Mg as well as
decrease the availability of P in root zone, which reflected on increase these nutrients in
leaves of cucumber plants.

3.1.3. Effect of microorganisms inoculation


Data in Table (46) declare that leaves of cucumber plants inoculated with Azotobacter
were significantly higher in contents of nitrogen (4.35 g/100g dry matter) and calcium (196
mg.kg-1), in the first season only, if compared to other inoculation treatments. However, it
could be observed that contents of potassium, calcium and magnesium were higher in the
second season if compared to the first season.
In addition, phosphorus us increased significantly in leaves of cucumber plants
inoculated with Glomus (0.51, 0.54 g/100g dry matter) or Glomus + Azospirillum (0.50, 0.51
g/100g dry matter) in the two growing seasons, respectively.
Meanwhile, inoculation of cucumber plants with Glomus + Azotobacter caused a
significant increase in potassium content (1440 mg.kg-1) comparing to control (1323 mg.kg-1)
in the first season, while no significant difference were found in the second season.
Table 46. Effect of microorganisms inoculation on N,P,K, Ca and Mg contents of
cucumbers leaves in 2006 and 2007

N P K Ca Mg
Treatments
g/100g dry matter mg.kg-1 fresh weight

Season 2006

N1 4,35 0,42 1378 196 122

N2 4,07 0,37 1358 183 127

N1+N2 4,19 0,43 1361 191 123

G 4,18 0,51 1372 179 114

G+N1 3,91 0,42 1440 174 111

G+N2 4,09 0,50 1335 187 128

G+N1+N2 4,01 0,41 1356 172 121

Control 4,22 0,37 1323 175 129

L.S.D. 0,25 0,05 63 12 11

Season 2007

N1 3,57 0,44 1858 242 136

N2 3,63 0,41 1829 233 145

N1+N2 3,54 0,45 1887 217 135

G 3,68 0,54 1853 256 135

G+N1 3,63 0,44 1762 233 144

G+N2 3,56 0,51 1874 241 135

G+N1+N2 3,63 0,42 1784 215 125

Control 3,68 0,40 1739 245 137

L.S.D. NS 0,04 NS NS 12
N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
Magnesium recorded significant decrease in leaves of cucumber plants when inoculated
with Glomus + Azotobacter (111 mg.kg-1) in the first season (129 mg.kg-1) than control (129
mg.kg-1).
On contrast, in the second season, inoculation cucumber plants with Azospirrillum
(145 mg.kg-1) or Glomus + Azotobacter (144 mg.kg-1) presented higher significant increase in
Mg content in leaves of cucumber than control (137 mg.kg-1).
However, these results are agreed with those obtained by Shahaby and Agwa (1993).
The authors found that inoculation seeds of spinach with Azotobacter chroococcum alone or
plus Azospirillum brasilense increased significantly total nitrogen, uptake of phosphorus us
and potassium in spinach leaves. Furthermore, inoculation of tomato with Azospirillum and
phosphate dissolving bacteria (PDB) stimulated a higher uptake of nitrogen and phosphorus
(Poi, 1998).
Similarly, Demir (2004) mentioned that leaves of pepper plants inoculated with Glomus
intraradices showed better phosphorus comparing to un-inoculated plants. These results are in
harmony with Neumann (2005) who reveled that AM fungal inoculation increased
phosphorus us uptake of tomato.
It can be suggested that PGPR improved mineral uptake due to increases in specific
ion fluxes at the root surface (Bertrand et al, 2000). On the other hand, Marchal and
Vanderleyden (2000) mentioned that excessive flux of O2 would inhibit nitrogenase activity
and supplying the fixed N2 to the plant.

3.1.4. Effect of interactions


Interaction between cultivar x substrate did not show any significant differences in
terms of nitrogen, phosphorus, potassium and magnesium in the first season, as shown in
Table (45).
Only calcium recorded significant increase in leaves of cultivar Passandra when grown
in rockwool (199 mg.kg-1).
In the second season, nitrogen (3.8 g/100g dry matter) and potassium (2061 mg.kg-1)
revealed significant increase in leaves of cultivar Passandra grown in peatmoss (Figs. 28-29).
Similar results were obtained with cultivar Girola grown in peatmoss in terms of calcium and
magnesium (283 and 148 mg.kg-1), as shown in Table (45). From the above results it could be
noticed that leaves of both cultivars showed significant increase of N, K, Ca and Mg when
grown under peatmoss if compared to rockwool in 2007.
Fig.28. Effect of interaction between cultivar x substrate on nitrogen
content in leaves of cucumbers in 2007

4,0

3,8
g/100g dry mater

3,6

3,4

3,2

3,0
Rockwool Peatmoss

Passandra Girola

Fig.29. Effect of interaction between cultivar x substrate on potassium


content in leaves of cucumbers in 2007

2250
mg.kg fresh weight

2000
-1

1750

1500
Rockwool Peatmoss

Passandra Girola

According to the interaction between cultivar x microorganisms inoculation, data in


Table (47) present that inoculation of cucumbers with Azotobacter increased significantly
nitrogen content in leaves of cultivar Girola (4.38 g/100g dry matter) and cultivar Passandra
(4.33 g/100g dry matter) if compared to other inoculation treatments in the first season only
Fig (30).
Phosphorus contents in Table (49) reveal non-significant increase in leaves of cultivar
Girola when inoculated with Glomus in both growing seasons (0.54 and 0.58 g/100g dry
matter) than control (0.36 and 0.40 g/100g dry matter), respectively.

Fig.30. Effect of interaction between cultivar x microorganisms inoculation


on nitrogen content in leaves of cucumbers in 2006

4,5

4,3
g/100d dry matter

4,0

3,8

3,5

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Also phosphorus in leaves of cultivar Passandra presented non-significant increase in


both growing seasons (0.51 and 0.50 g/100g dry matter) when inoculated with Glomus +
Azospirillum, as shown in Figs (31-32).
In Table (51), potassium content in leaves of cultivar Girola showed the highest value
with Glomus + Azotobacter inoculation (1438 mg.kg-1), and cultivar Passandra with Glomus
(1458 mg.kg-1) inoculation, in the first season if compared to control. Meanwhile, in the
second season, the highest level of potassium in leaves of cultivar Girola (1892 mg.kg-1)
found when plants inoculated with Glomus + Azospirillum, and (1997 mg.kg-1) in cultivar
Passandra when plants inoculated with Azotobacter + Azospirillum.
However, these influence of microorganisms inoculation treatments on contents of
potassium and phosphorus did not reach to significant level, but positively increased contents
of both nutrients in leaves in the two cultivars.
Fig.31. Effect of microorganisms inoculation on phosphorus content in leaves
of cultivar Girola 2006 and 2007

0,70

0,60
g.100g dry matter

0,50

0,40

0,30

l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig.32. Effect of microorganisms inoculation on phosphorus content in leaves


of cultivar Passandra 2006 and 2007

0,60
g.100g dry matter

0,50

0,40

0,30
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Calcium in Table (53), showed significant increase (247 mg.kg-1) in leaves of cultivar
Girola in the second season with Azotobacter inoculation treatment, while cultivar Passandra
reveal the highest significant increase (263 mg.kg-1) when inoculated with Glomus +
Azotobacter or Glomus + Azospirillum (Fig. 33).

Fig.33. Effect of interaction between cultivar x microorganisms inoculation


on calcium content in leaves of cucumbers in 2007

300
mg.kg fresh weight

250

200
-1

150

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Magnesium did not show significant differences with inoculation treatments in both
growing seasons if compared to control, as shown in Table (55).
As regard to the effect of interaction between substrate x microorganisms inoculation,
data in Table (48) show that nitrogen content in leaves of cucumbers recorded significant
increase (4.74 g/100g dry matter) when plants grown in rockwool and inoculated with
Azospirillum comparing to other treatments. In general, content of nitrogen were higher in
leaves of cucumber plants when grown in rockwool than peatmoss in the first season, while
no differences were observed in the second season.
Furthermore, phosphorus did not reveal significant differences between treatments in
both growing seasons Table (50).
In addition, potassium content in leaves of plants inoculated with Glomus recorded the
best significant increase (1448 mg.kg-1) when grown in peatmoss than rockwool (1398 mg.kg-
1
) in the first season only, as shown in Fig. (34).
Moreover, in the first season, calcium in leaves of cucumbers showed higher
significant values (206, 205 and 204 mg.kg-1) when plants grown in peatmoss and inoculated
with Azospirillum, Azotobacter or Glomus, respectively, as shown in Fig (35). In the second
season, contentss of calcium showed the best significant increase with Glomus + Azotobacter
or Glomus alone (309 and 303 mg.kg-1), respectively.

Fig.34. Effect of interaction between substrate x microorganisms inoculation


on potassium content in leaves of cucumbers in 2006

1500

1400
mg.kg fresh weight

1300
-1

1200

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Rockwool Peatmoss

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig.35. Effect of interaction between peatmoss x microorganisms inoculation


on calcium content in leaves of cucumbers in 2006 and 2007

400
mg.kg fresh weight

300

200
-1

100
l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


On the contrary, content of magnesium in leaves of cucumbers recorded the best
significant increase in the first season only with plants grown in rockwool and inoculated with
Glomus (141 mg.kg-1) or Glomus + Azospirillum (140 mg.kg-1) comparing to other treatments
(Table 56).
The above results indicate that microorgismsm inoculation affected positively
nutrients contents in leaves of cucumber plants. This influence was varied according to the
type of substrates. It is recognized that the bacterium’s growth promoting effects originate
from mechanisms other than N2 fixation (Dobbelaere et al, 2001).
Therefore, the most common method of PGPR stimulation of plant growth is actually
production of phytohormones which increase root surface area; hence turn enables the plant to
absorb more nutrients to enhance growth of the entire. Barber and Cushman (1981), reported
that nutrient uptake in plants indicate the extremely important role of root surface area as a
factor influencing the capacity of plants to absorb inorganic nutrients.
The effect of phytohormone productivity by PGPR on host is evident in a number of
root parameters, namely greater root mass, more root branching, thinner roots, and more root
hair. Furthermore, PGPR are common to increase root mass (Vessey and Buss, 2002).
Molla et al (2001) showed that Azospirrilium braselinse Sp7 caused a 63% increase in
soybean root dry weight more than six fold increase in specific root length, and more than a
ten fold increase in total root length.
It can be concluded that positive effects of PGPR on roots of cucumber plants
increased their ability to absorb more nutrients. However, it’s well known that, under soilless
condition, rockwool substrate has optimal aeration and hold a large amount of solution, than
peatmoss, which facilitate function of cucumber root to uptake nutrients.
IV. 3.2. Mineral composition of fruits
3.2.1. Effect of cultivar
Results in Table (57) declare that nitrogen content showed higher significant increase
in fruits of cultivar Passandra (2.88 g/100g dry matter) in the first season and with cultivar
Girola (3.40 g/100g dry matter) in the second one.
On contrast, contents of calcium, phosphorus and potassium in the first season
presented significant increase in fruits of cultivar Girola than fruits of cultivar Passandra.
However, phosphorus and potassium contents in fruits of both cultivars did not reveal
significant differences in the second season. In this respect, it can be suggested that these
differences in nutrient contents between both cultivars could be related to the effect of
genotype.

3.2.2. Effect of substrate


Generally, fruits of cucumbers grown in rockwool or peatmoss did not show
significant differences in content of tested nutrients in both growing seasons with some
exceptions, as shown in Table (57). These findings agree with Schroeder and Sell (2009) who
mentioned that nutrient content of cucumber fruits did not differ significantly between the
substrates.
In the first season, nitrogen and magnesium reveal significant increase when grown in
rockwool (2.87 g/100g dry matter and 111 mg.kg-1) than peatmoss (2.81 g/100g dry matter
and 95 mg.kg-1), respectively. This might be attributed to the high concentration of those
nutrients in leaves of cucumber plants in the first season. These results were confirmed by
Table (45) which reveals significant increase of nitrogen (4.45 g/100g dry matter) and
magnesium (136 mg.kg-1) in leaves of cucumber plants when grown under rockwool, in the
first season.
However, these differences in macro and micronutrients could be explained based on
different response of both substrates to the used nutrient solution. Similar suggestion was
mentioned by Colla et al (2003).
On contrast, potassium and calcium contents in fruits of cucumbers were significantly
higher under peatmoss (1292 and 235 mg.kg-1) if compared to rockwool (1195 and 164
mg.kg-1) in the second season. In accordance, Gomez et al (2002 b) declares that potassium
concentration was higher in coir than rockwool as a growing media, while magnesium
concentration was not significantly affected by the type of substrate.
In this respect, it can be suggested that this increase in K and Ca contents could be
related to the high uptake of those nutrients under peat than rockwool. In connection,
Cantliffe et al (2003) and Iapichino et al (2007) mentioned that non-mineral medias such as
pine bark, coconut coir and sphagnum peat mixtures became more used for commercial
production, because of its satisfactory aeration and water holding capacity than inert
substrates. In addition, high loss of water and nutrient elements from rockwool slabs caused a
decrease in nutrient absorption by plant root (Benton, 2005).

3.2.3. Effect of microorganisms inoculation


Table (58) declares that nitrogen and phosphorus content in fruits of cucumbers were
significantly higher under most of microorganisms inoculation treatments if compared to
untreated treatment in both growing seasons.
Similarly, potassium increased significantly under microorganisms inoculation in the
first season and recorded the highest content in fruits of cucumbers (1087 mg.kg-1) when
plants inculcated with Azotobacter + Azospirillum than control (935 mg.kg-1).
Meanwhile, calcium presented the highest content (202 mg.kg-1) with Azospirillum
inoculation, and magnesium (109 mg.kg-1) with Azotobacter inoculation if compared to
control (176 and 98 mg.kg-1), respectively. On contrast, differences between inoculated and
control plants did not reach to significant level in the second season.
In connection, Garib (2002) found that nitrogen and dry matter content in cucumbers
leaves were increased using combination of Azospirillum + Azotobacter + phosphate
dissolving bacteria. In addition, concentrations of nitrogen and phosphorus us in roots and
magnesium, copper, and zinc concentration in shoots of cucumber were increased by Glomus
mosseae inoculation (Changxian Wang et al, 2006).
Furthermore, microorganisms produce an IAA and cytokinin (Rai, 2005). IAA causes
an increase in root length (Patten and Glick, 2002). Cytokinins promote cell divisions, cell
enlargement, and tissues expansion in certain plant parts (Salisbury, 1994).
In this respect, it could be said that Azotobacter and/or Azospirillum inoculation
improved nutrients content in cucumber fruits if compared to the control. This conclusion
based on fact that microorganisms play a vital role to increase nutrients contents and plant
metabolism processes in leaves and shoots of inoculated plants.
Table 58. Effect of microorganisms inoculation on N,P,K, Ca and
Mg contents of cucumber fruits in 2006 and 2007

N P K Ca Mg
Treatments
g/100g dry matter mg.kg-1 fresh weight

Season 2006

N1 2,87 0,33 1029 174 109

N2 2,83 0,33 1075 202 102

N1+N2 2,62 0,33 1087 187 105

G 2,92 0,32 1054 177 105

G+N1 3,03 0,32 1061 167 106

G+N2 2,79 0,33 1053 169 102

G+N1+N2 2,78 0,32 1041 178 100

Control 2,88 0,27 935 176 98

L.S.D. 0,11 0,03 65 20 6

Season 2007

N1 3,30 0,38 1277 200 83

N2 3,28 0,40 1207 178 84

N1+N2 3,29 0,38 1261 222 84

G 3,18 0,42 1236 244 85

G+N1 3,19 0,44 1270 169 83

G+N2 3,30 0,40 1294 193 79

G+N1+N2 3,23 0,45 1178 209 83

Control 2,70 0,38 1225 184 85

L.S.D. 0,33 0,04 NS NS NS


N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
3.2.4. Effect of interaction
The interaction between cultivar x substrate did not affect contents of nitrogen,
phosphorus and magnesium in both growing seasons. Meanwhile, potassium content in fruits
of cultivar Girola showed the highest content (1402 mg.kg-1) when grown under rockwool in
the first season, as shown in Table (57).
On the contrary, content of calcium in fruits of cultivar Girola grown in peatmoss
recorded the highest significant increase (199 mg.kg-1) in the first season and in fruits of
cultivar Passandra (275 mg.kg-1) in the second season.
Concerning the effect of interaction between cultivar x microorganisms inoculation,
data in Table (59) show that nitrogen content in fruits of cultivar Passandra was significantly
higher (3.18 g/100g dry matter) with Glomus + Azotobacter followed by Azospirillum (2.99
g/100g dry matter) in the first season. In contrast, second season reveals significant increase
in fruits of cultivar Girola (3.75 and 3.61 g/100g dry matter) when cucumber plants inoculated
with Azotobacter + Azospirillum or Glomus + Azospirillum, respectively, as shown in Figs
(36-37).

Fig.36. Effect of interaction between cultivar x microorganisms inoculation


on nitrogen content in fruits of cucumbers in 2006

3,5
g/100g dry matter

3,0

2,5

2,0
l
G

2
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


Fig.37. Effect of interaction between cultivar x microorganisms inoculation
on nitrogen content in fruits of cucumbers in 2007

4,0
g/100g dry matter

3,5

3,0

2,5

2,0

l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Furthermore in Table (61), phosphorus content in fruits of cultivar Girola presented


higher significant increase (0.38 g/100g dry matter) with Azotobacter + Azospirillum
inoculation in the first season, and (0.48 and 0.45 g/100g dry matter) with Glomus and
Glomus + Azotobacter + Azospirillum, respectively, in the second season, if compared to
other treatments (Fig. 38).

Fig.38. Effect of microorganisms inoculation on phosphorus content


in fruits of cultivar Girola in 2006 and 2007

0,60

0,50
g/100g dry matter

0,40

0,30

0,20
l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


Moreover, calcium content in fruits of both cultivars did not show significant
differences in both growing seasons (Table 65).
On the other hand, potassium in the first season (Table 63) and magnesium (Table 67),
recorded higher significant increase (1146 and 109 mg.kg-1), respectively, in fruits of cultivar
Girola when inoculated with Glomus + Azotobacter + Azospirillum, as shown in Figs (39-40).

Fig.39. Effect of interaction between cultivar x microorganisms inoculation


on potassium content in fruits of cucumbers in 2006

1200

1100
mg.kg fresh weight

1000
-1

900

800

l
1

2
1

G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig.40. Effect of interaction between cultivar x microorganisms inoculation


on magnesium content in fruits of cucumbers in 2006

100

90
mg.kg fresh weight

80

70
-1

60
l
1

2
1

G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


However, it can be observed that fruit of both cucumber cultivars presented higher
contents of mineral nutrients than control, in the second season, with all microorganisms
inoculation.
The interaction between substrate x microorganisms inoculation did not present
significant differences in contents of nitrogen, potassium and calcium in both seasons (Tables
60, 64 and 66).
In addition, Table (62) shows that phosphorus content in fruits of cucumbers
presented higher significant increase (0.51 g/100g dry matter) when grown in rockwool or
peatmoss inoculated with Glomus + Azotobacter + Azospirillum in the second season only
(Fig. 41).

Fig.41. Effect of interaction between cultivar x microorganisms inoculation


on phosphorus content in fruits of cucumbers in 2007

0,6
g.100g dry matter

0,5

0,4

0,3
l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

Peatmoss Rockwool

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Meanwhile, fruits of cucumbers grown in rockwool and inculcated with Azotobacter


reveal in Table (68) higher significant increase of magnesium (118 mg.kg-1) if compare to
other treatment in the first season (Fig. 42).

The above results reveal that fruits of cultivar Girola showed better nutritional
contents than cultivar Passandra under the microorganisms inoculation treatments, as shown
in Figs (37, 38, 39 and 40). In addition, it can be observed that type of substrate, in general,
inhibited both of microorganisms and cultivars to express their interaction. In this respect, it
can be concluded that type of substrate is a critical factor affected nutritional status of
cucumber plants under the experiment conditions.
Fig.42. Effect of interaction between rockwool x microorganisms inoculation
on magnesium content in fruits of cucumbers in 2006.

120
mg.kg fresh weight

115

110
-1

105

100

l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

These could be related to the diurnal changes in pH or salinity of the nutrient solution
in growing media. In this respect, it can be suggested that many reasons could cause these
changes in composition of nutrient solution i.e., the low irradiance under greenhouse
condition reflected negatively on growth rate of greenhouse plants and leads to a lower
nutrient absorption and utilization. Also, due to low relative humidity under air-conditioning,
the relative rate water loss would normally be higher than the nutrient uptake, and this may
result in an overall increase in the nutrient concentration, thus creating salinity problem
(Awang et al, 2007).
IV. 4. Chemical quality
IV. 4.1. Vitamin C and total soluble solid (TSS)
4.1.1. Effect of cultivar
Data in Table (69) indicate that vitamin C and total soluble solid did not differ between
the two cucumber cultivars in both growing season, except cultivar Passandra which showed
significant increase in TSS (4.1 %) if compared to Girola cultivar (3.9 %) in the first season.
However, it can be noticed low variations between both cultivars in terms of vitamin C
and TSS values. In this respect, it can be concluded that type of cultivar did not affect vitamin
C or TSS contents. Similar suggestion was obtained by Hanafy Ahmed et al (2004) on sweet
pepper.

4.1.2. Effect of substrate


Type of substrate did not affect vitamin C in both growing seasons and TSS in the first
season, as shown in Table (69). On contrast, total soluble solid in the second season presented
significant increase in fruits of cucumber grown in peatmoss (4.5 %) than rockwool (3.4 %).
These results agreed with Colla et al (2003) who found no differences in fruit quality
between cucumbers grown in rockwool, perlite and coir. Also, Guler et al (1995) did not
present differences in soluble solid content among melon fruits grown in rockwool, perlite,
coir slabs or coir dust + perlite mixture. In connection, Peyvast et al (2008) did not indicate
significant effects on soluble solid in fruits of cucumber grown in peat, perlite or rice-hull
substrates.
On the other hand, Tuzel et al (2001) recorded significant increases in vitamin C and
total soluble solid of tomato fruits when grown in volcanic ash compared to perlit substrates.
Similarly, Inden and Torres (2004) mentioned that soluble solids content was higher in plants
grown in coconut coir (7.3 o Brix ) followed by rockwool (6.6 o Brix) and perlite (5.6 o Brix)

4.1.3. Effect of microorganisms inoculation


Inoculation cucumber plants with Glomus + Azospirillum recorded, in Table (70), the
highest significant increase in vitamin C (138 and 149 mg.kg-1) in both seasons, respectively,
Fig (43).
Also, total soluble solid revealed similar trend in the first season and recorded the
highest value (4.1 %) when inoculated with Glomus + Azotobacter + Azospirillum, as shown
in Fig. (44). No differences between inoculation treatments were found in the second season.
Table 70. Effect of microorganisms inoculation on fruit duality of cucumber
fruits in 2006 and 2007

Vitamin C TSS DM NO3- Sugars


Treatments
mg.kg-1 % % mg.kg-1 mg.kg-1

Season 2006

N1 124 4,1 2,2 158 16,8

N2 126 4,1 2,1 168 16,4

N1+N2 126 3,9 2,3 154 16,1

G 130 4,0 2,3 139 18,0

G+N1 120 4,0 2,3 115 16,6

G+N2 138 4,1 2,2 113 20,4

G+N1+N2 128 4,1 2,2 111 21,8

Control 114 4,1 2,1 96 19,3

L.S.D. 8,0 0,16 NS 21 NS

Season 2007

N1 137 4,0 2,9 335 18,3

N2 125 4,0 3,0 250 17,4

N1+N2 137 4,0 2,9 354 16,9

G 143 4,0 3,1 302 16,3

G+N1 135 3,8 2,9 309 17,9

G+N2 149 3,9 3,0 296 19,2

G+N1+N2 130 3,8 3,1 288 18,1

Control 115 4,1 3,3 304 19,1

L.S.D. 21 NS 0,24 NS 2,4


N1= Azotobacter chroocooccum N2= Azospirillum brasilense G= Glomus mosseae
Fig. 43. Effect of microorganisms inoculation on vitamin C content in fruits of
cucumbers in 2006 and 2007

160

140
mg.kg fresh weight

120
-1

100

nt
G

2
1

2
1

N2

+N

+N

N
N

co
1+
1+

+N
N

2006 2007 G

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig.44. Effect of microorganisms inoculation on TSS content


in fruits of cucumbers in 2006 and 2007

4,50

4,25

4,00
%

3,75

3,50
l
1

2
G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae


Siqueira and Saggin Junior (1995) mentioned that the most important benefit of
mycorrhiza is the increase in nutrient absorption, such as H2PO4-, Zn++, Cu++ and NH4+. These
benefits lead to improve the nutritional status of the plants, plant growth (Rai, 2005) and plant
water relationships (Schwab et al, 1991).
In connection, Qi GuoHui et al (2001) found that AM fungi Glomus intraradices,
Glomus mosseae, Glomus versiforme increased ascorbic acid in strawberry fruits. Similarly,
Lu GuiYun et al (2006) reported that same AM fungi significantly improved the quality of
cucumber fruits. In addition, mycorrhizal plants can enhance the accumulation of amino acids
(Johansen et al, 1996) and cause alternations in the content of hormones (Goicoecheta et al,
1997), and increase enzymatic activity (Ruiz-Lozano and Azcon, 1996), as well as secondary
metabolites (Peipp et al, 1997).
However, the above results declare the positive effects of single inoculation of N-
Fixers, and the combined microorganisms Glomus + Azospirillum or Glomus + Azotobacter +
Azospirillum inoculation on quality of cucumber fruits.

4.1.4. Effect of interaction


The interaction between cultivar x substrate showed the highest content of vitamin C
in fruits of cultivar Passandra grown in rockwool in both growing seasons (132 and 142
mg.kg-1), respectively.
These results in Table (69) do not agree with many investigations indicated that
organic growing medias are more efficiency to improve vitamin C content in cucumber fruits.
Tuzel et al (2001) found significant increases in vitamin C of tomato fruits when grown in
volcanic ash compared to perlite substrates. Also, fruits of cherry tomato grown on coco-fiber
contained much more vitamin C than those grown on rockwool (Holtmann and Kobryn 2003).
In this respect, it can be suggested that these unexpected results could be due to the sampling
procedure (Bohme et al, 2008).
On contrast, total soluble solid did not differ under both types of growing media.
Similar findings were mentioned by Guler et al (1995) on melon, Maniutiu et al (1995) on
cucumber and Shinohara et al (1999) on tomato.
According to the effect of cultivar x microorganisms inoculation interactions, Fig. (45)
declare that vitamin C in fruits of cultivar Passandra (139 mg.kg-1) and cultivar Girola (138
mg.kg-1), in the first season, were significantly higher when cucumbers inoculated with
Glomus + Azospirillum if compared to other inoculation treatments (Table 71). No significant
results were found in the second season.
On the other hand, un-inoculated plants of cultivar Passandra showed in Table (73) the
highest non-significant value of total soluble solid (4.4 %), in the second season if compared
to inoculated plants (Fig 46).

Fig.45. Effect of interaction between cultivar x microorganisms inoculation


on vitamin C content in fruits of cucumbers in 2006

150

130
-1
mg.kg

110

90

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Fig.46. Effect of interaction between cultivar Passandra x microorganisms


inoculation on TSS in fruits of cucumbers in 2006

5,0

4,5

4,0
%

3,5

3,0
l
1

2
1

G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

Meanwhile, Table (72) show that interactions between substrate x microorganisms


inoculation reveal significant increases in vitamin C of cucumbers grown under peatmoss
(143 mg.kg-1) or rockwool (134 mg.kg-1) and inoculated with Glomus + Azospirillum in the
first season only (Fig. 47).

Fig.47. Effect of interaction between substrate x microorganisms inoculation


on Vitamin C content in fruits of cucumbers in 2006

160

140
-1

120
mg.kg

100

80

l
1

2
1

G
N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
Peatmoss Rockwool

N1= Azotobacter chroococcum G


N2= Azospirillum brasilense G= Glomus mosseae

Table (74) declares that total soluble solid presented the best significant increase (4.3
%) with cucumbers grown in peatmoss and inoculated with Glomus + Azotobacter +
Azospirillum, Fig (48). On the other hand, no differences in vitamin C and TSS were observed
in the second season.
These results show that mixture of Glomus and Azospirillum inoculation under
peatmoss substrate led to improve quality of cucumbers. These positive influences can be
explained according to the role of mycorrhizal fungi to cause useful physiological alterations
such as increasing the stomatal conductance, respiration, transpiration, absorption of CO2,
photosynthesis (Mathus and Vyas, 1995; Ruiz-Lozano et al, 1995; Goicoecheta et al, 1997;
Rilling et al, 2002). Meanwhile, Azospirillum is able to produce plant hormones as well as
polyamines and amino acids in culture (Thuler et al, 2003). So, it can be suggested that dual
inoculation combinations (bacterium - fungus) have proven beneficial in plant growth
promotion than single strains alone (Park et al, 1988).
Schroth and Becker (1990) mentioned that mixtures of microorganisms might be
more efficacious than single strains in promoting plant growth. Therefore, Srinath et al, 2003
mentioned that application of Azospirillum as PGPR have been found to stimulate the level of
infection of AM of the Glomus genus. In accordance, Shabayev et al (1996) found that
photosynthesis in soybean inoculated with mixed cultures of Bradyrhizobium japonicum and
Pseudomonas fluorescens as well as Glomus mosseae were due to growth promoting
substances.

Fig.48. Effect of interaction between peatmoss x microorganisms inoculation


on TSS in fruits of cucumbers in 2006 and 2007

5,0

4,5

4,0
%

3,5

3,0

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
2006 2007

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

In this respect, it can be concluded that type of cultivar or substrate, as single factor,
did not affect vitamin C or TSS content in fruits of cucumbers in both growing seasons with
some exceptions. In addition, inoculation of cucumber plants with Glomus + Azospirillum
improved vitamin C contents in fruits of both cucumbers cultivars, especially when
cucumbers grown in peatmoss substrate. Similarly, TSS contents were significantly higher in
fruits of cucumbers grown in peatmoss, but when inoculated with Glomus + Azotobacter +
Azospirillum.
IV. 4.2. Dry matter content, nitrate and total sugars concentrations
4.2.1. Effect of cultivar
Concerning the effect of cultivar on dry matter content, nitrate and total sugars
concentrations of cucumber fruits, data in Table (69) clear that dry matter of cultivar
Passandra fruits were significantly higher (2.3 and 3.1 %) than cultivar Girola (2.1 and 2.9 %)
in both seasons, respectively.
Total sugars showed similar trend in the second season with cultivar Passandra (19.6
mg.kg ) if compared to cultivar Girola (16.8 mg.kg-1), while no difference were recorded in
-1

the first season between both cultivars. On contrast, fruits of cultivar Girola presented higher
significant increase of nitrate (145 and 351 mg.kg-1) than cultivar Passandra (117 and 258
mg.kg-1) in the two growing seasons, respectively.
In this respect, it can be suggested that mini fruit type of cucumbers such as cultivar
Passandra, are more active to absorb more nutrients, hence rate of translocation and
accumulation of such nutrients in tissue of mini fruit type is higher than long fruit type, as
cultivar Girola.

4.2.2. Effect of substrate


Fruits of cucumbers grown in rockwool showed significant increase in dry matter and
nitrate concentration (3.5 % and 342 mg.kg-1) in the first season only if compared to those
grown in peatmoss, as shown in Table (69). Both growing media did not influence dry matter
or nitrate accumulation in the second season. In this respect, it can be suggested that the low
anion holding capacities of peat lead to decrease of nutrients absorption in root zone and
increase the need for frequent fertilizer application in greenhouse culture (Rai, 2005).
In connection, Martyn (1996) and Bohme et al (2006) mentioned that rockwool is the
most suitable inert substrate in soilless systems because of its ability to hold large amounts of
nutrient solution and has optimal aeration.
On contrast, total sugars concentration did not differ significantly between substrates
in both growing seasons. In accordance, Colla et al (2003) revealed no significant differences
in fruit quality between cucumbers grown in rockwool, perlite and coir. Similarly, Peyvast et
al (2008) found no significant effects on soluble solid in cucumber grown in peat, perlite or
rice-hull substrates.
4.2.3. Effect of microorganisms inoculation
Data in Table (70) show that dry matter and total sugars contents in the first season as
well as concentration of nitrate in the second one did not was influenced by any of
microorganisms inoculation treatments. These results are in harmony with Gul et al (2007).
The authors found that inoculation of tomato with Bacillus subtilis or Trichoderma horzianum
did not affect fruit quality characteristics.
However, significant decrease in dry matter was observed with Azotobacter (2.9 %)
and Azospirillum (3.0 %) if compared to control (3.3 %) in the second season.
On the other hand, nitrate concentrations were significantly increased with
Azotobacter (158 mg.kg-1), Azospirillum (168 mg.kg-1) if compared to Glomus (139 mg.kg-1)
or control (96 mg.kg-1) in the first season. Similar trend were found in the second season in
spite of non-significant differences.
These results declare the role of PGPR in fixing atmospheric nitrogen that is
transferred to the plant and make it available to the plant root, solubilization of minerals such
as phosphorus us, and synthesis of phytohormones (Glick, 1995; Dobbelaere et al, 2001).
In connection, Ribaudo et al (2001) pointed out to the biological fixation role of
Azospirillum sp. in fixing nitrogen and increase the activity of glutamate dehydrogenase and
glutamine synthetase.
In this respect, Hanafy Ahmed et al (2002 b) pointed out that N supply is one of the
most important factors affecting nitrate accumulation in growing plants.
Moreover, total sugars presented opposite trend and increased significantly with
control treatment (19.1 mg.kg-1) than Azotobacter (18.3 mg.kg-1), Azospirillum (17.6 mg.kg-1)
or their mixture (16.9 mg.kg-1). Meanwhile, the highest significant increases were recorded
with Glomus + Azotobacter (19.2 mg.kg-1), in the first season.
However, the above results declare the negative relationship between nitrate
accumulation and total sugars concentration in fruits of cucumbers.
Similar results were obtained by Hanafy Ahmed et al (1997) on Jew’s mallow and
radish plants and Hanafy Ahmed et al (2002 a,c) on sweet pepper and rocket plants,
respectively.
Furthermore, it is important here to mention that increasing the level of N and P
increased both fresh and dry weights of cucumber leaves more than control. Therefore, these
decreases in total sugar concentrations which were concomitantance with increases in the
fresh and dry weights of cucumber plants may be due to that the plants used most of carbon in
tructural growth, but incorporated relatively less carbon in soluble organic compounds.
Similar results and suggestions were obtained by Hanafy Ahmed et al (2002 b) and (2004) on
lettuce and sweet pepper plants, respectively.
In this respect, it can be suggested that, nitrate accumulation might be increased in the
vacuoles to compensate the shortage of sugars. So, it can be concluded that NO-3
accumulation was inversely related to accumulation of sugars. Similar findings were reported
by Blom-Zandstra and Lampe (1985) and Hanafy Ahmed (1996) on lettuce, Hanafy Ahmed et
al (1997) on Jew’s mallow and radish as well as Hanafy Ahmed et al (2002 a, b and c) on
rocket, lettuce and sweet pepper, respectively. In this respect, Blom-Zandstra et al (1988)
mentioned that the lettuce genotype which had higher nitrate concentration had also lower
concentration of sugars and organic compounds.
In addition, the increases in the concentration of total sugar may also be explained on
the assumption that, as already known, the nitrate accumulation in the vacuoles is not readily
available. However, when the nitrogen is not sufficient amount, the nitrate might remove from
the vacuoles to sustain the protein synthesis and the plant will store sugars and organic acids
to compensate for the declining osmotic value. This suggestion is in a good agreement with
that of Blom-Zandstra (1989).

4.2.4. Effect of interaction


Table (69) shows that values of dry matter, nitrate and total sugar contents in fruits of
cultivar Passandra or Girola did not differ significantly under both substrates in the two
growing season, with some exceptions in the second season. Similar results were obtained by
Guler et al and Maniutiu et al (1995), Shinohara et al (1999) as well as Nurznynski (2006).
Cultivar Passandra expressed these exceptions and recorded the highest significant
increases in dry matter (3.7 %) when plants grown in rockwool, while total sugars reveal
higher significant increase (19.8 mg.kg-1) under peatmoss.
Moreover, other interactions treatments did not show significant differences, as shown
in Tables (75-80) in both growing seasons.
The only exception was found in Table (79) which showed that fruits of cultivar
Passandra and Girola revealed higher significant increase in total sugars (25.6 mg.kg-1) when
inoculated with Glomus + Azotobacter + Azospirillum, if compared to other inoculation
treatments in the second season (Fig. 49). Generally, it can be suggested that both growing
media did not interact positively with microorganisms inoculation.
Fig.49. Effect of interaction between cultivar x microorganisms inoculation
on total sugars content in fruits of cucumbers in 2007

30

25
-1
mg.kg

20

15

10

l
1

2
G
1

N2

ro
+N

+N
N

nt
1+

1+

Co
G

G
N

+N
G
Passandra Girola

N1= Azotobacter chroococcum N2= Azospirillum brasilense G= Glomus mosseae

In accordance, Vestberg and Kukkonen (2009) mentioned that peat growing media
have high populations of fungi and bacteria, in particular actinomycetes, which may affect
symbiosis efficiency of AM. In addition, extra cellular enzymes like ß-glucosidase,
phosphatase, sulphatase, xylosidase, esterase present in peat may negatively interact with
microorganisms population (Freeman et al, 1995).
V. SUMMARY
This experiment was carried out during the two successive seasons of 2006 and 2007
under the open soilless system in greenhouse at Faculty of Horticulture in Lednice, Mendel
University, Czech Republic. Transplants of cucumber (Cucumis sativus L. cultivars
‘Passandra’ and ‘Girola’) were arranged in 4 replicates using split-split plot design. Each
cultivar was grown on rockwool and peatmoss as based growing media. Transplants were
inoculated by seven microorganisms treatments: N1 (Azotobacter chroococcum), N2
(Azospirillum brasilense), G (Glomus mosseae), and their combinations N1 + N2, G + N1, G
+ N2, G + N1 + N2 and control.
Generally, it can be observed that cultivar Passandra was exceeded cultivar Girola in
terms of plant height, number of leaves as well as leaf FW and leaf DW. On contrast, leaf area
of cultivar Girola showed higher significant increase in the 2nd season. Early and total yield of
both cultivars did not reveal much variation. These results indicate that each cultivar
expressed its effect on vegetative growth characteristics based on variation among both geno-
types. However, cultivar Girola presented higher significant increase in fruit length, diameter
and weight. Meanwhile, cultivar Passandra produced higher significant number of fruits/plant
in both seasons; combined with higher significant increase of grade “Extra” fruits were
obtained in the 2nd season.
Nitrogen, phosphorus and magnesium did not differ in leaves of both cultivars.
Moreover, K was significantly higher in leaves of cultivar Girola and Passandra in 1st and 2nd
seasons, respectively. Nitrogen in fruits of cultivar Passandra and P, K, Ca in fruits of cultivar
Girola revealed higher significant increases in the first season while N, P, and Mg did not
differ in leaves of the two cultivars, with some exceptions. Type of cultivar did not affect
vitamin C or TSS contents. Meanwhile, dry matter in fruits of cultivar Passandra were
significantly higher in both seasons; and total sugars in the 1st season. Nitrate, recorded higher
significant increase in fruits of cultivar Girola, but values still under the critical limits. In
general, it can be concluded that variations between cultivars are correlated to the gene-type,
and cultivars are varied in their yield productivity and quality.
According to the effect of substrate type on cucumber plants, data presented that plants
grown on peatmoss revealed significant increase in number of leaves/plant and leaf DW. Leaf
area and leaf FW showed different trends in both growing seasons. However, light significant
increase was obtained in early yield of plants grown on rockwool. On contrast, total yield of
cucumbers grown in peatmoss recorded higher significant increase in the 1st season. Similar
trend was noticed with plants grown on rockwool in the 2nd season.
Fruits of cucumbers grown in rockwool revealed lightly significant increase in fruit
length and weight. Meanwhile, cucumbers grown on peatmoss produced higher significant
fruits number/plant and marketable yield of grade “Extra” in the 1st season and when grown
on rockwool in the 2nd season. Macronutrients in leaves and fruits as well as vitamin C, total
sugars and TSS in fruits of cucumbers did not differ between substrates, with some
exceptions. Fruits of cucumbers grown in rockwool showed significant increase in dry matter
and NO3- in the 1st season. In this respect, it can be concluded that distinction in chemical and
physical properties between rockwool and peatmoss reflected on recorded results.
Cucumber plants inoculated by N-Fixing bacteria and/or Glomus did not affect growth,
early yield, fruit length, diameter and fruits number/plant in both seasons, with some
exceptions. However, it can’t be denied that microorganism inoculation treatments did light
positive effect on some of vegetative growth characteristics and yield component. Limited
light quantities inside greenhouse in winter season and/or the variation in pH and saltation of
the nutrient solution could inhibit the microorganisms formation under such conditions.
Furthermore, all inoculation treatment revealed better, but not significant, total yield.
These improve in total yield reached 8 and 21 % in the 1st and 2nd seasons, respectively. In
addition, significant increase in average fruit weight was found when cucumber plants
inoculated with Azotobacter alone or combined with Azospirillum than control in the 1st
season. Inoculating cucumber plants improved the marketable yield, contents of
macronutrients in leaves and fruits of both cultivars, with some exceptions.
Vitamin C in fruits of cucumber plants recorded the highest significant increase when
inoculated with Glomus + Azospirillum. In addition, total soluble solid revealed similar trend
in the 1st season when plants inoculated with Glomus + Azotobacter + Azospirillum.
Moreover, significant decrease in dry matter was observed with Azotobacter and Azospirillum
inoculation if compared to control in the 2nd season. Nitrate concentrations were significantly
increased with Azotobacter or Azospirillum treatments if compared to Glomus or control in
the 1st season.
Interactions between cultivars and substrate or microorganisms show preferable effect
on most of growth characteristics of cucumber plants. Particularly, N-Fixers alone/or
combined with Glomus improved growth of cucumber plants grown in peatmoss. Cucumber
plants grown in peatmoss and inoculated with Glomus + Azotobacter or Azospirillum, were
significantly higher in total yield in the 1st season than control. In general, it can be concluded
that growth and yield of cultivar Passandra grown in peatmoss were positively responded to
microorganisms inoculation than other treatments.
Leaves of both cultivars showed significant increase of N, K, Ca and Mg when grown
on peatmoss in 2nd season. Neither cultivar nor substrate as single factors affected vitamin C
or TSS content, with some exceptions. Dry matter, nitrate and total sugar in fruits of cultivars
did not differ under both substrates, with some exceptions in 2nd season. However, inoculation
of plants with Glomus + Azospirillum improved vitamin C contents in fruits of both cultivars,
especially when cucumbers grown on peatmoss. Similarly, TSS content was significantly
higher in fruits of cucumbers grown in peatmoss, but when inoculated with Glomus +
Azotobacter + Azospirillum. In this respect, it could be said that microorganisms inoculation
improved nutrients content in cucumber fruits if compared to the control. This conclusion
based on fact that microorganisms play a vital role to increase nutrients contents and plant
metabolism processes in leaves and shoots of inoculated plants.
V. ZÁVĚR
Práce probíhala v letech 2006 a 2007 v otevřeném hydroponickém systému ve
skleníku Zahradnické fakulty v Lednici, Mendelovy Univerzity v Brně, Česká Republika.
Rostliny okurek (Cucumis sativus L. odrůdy ‘Passandra’ a ‘Girola’) byly vysázeny ve 4
opakováních ve split-plot designu. Rostliny byly pěstovány v čedičové plsti a v rašelině.
Sadba byla inokulována těmito mikroorganismy: N1 (Azotobacter chroococcum), N2
(Azospirillum brasilense), G (Glomus mosseae), a jejich kombinace N1 + N2, G + N1, G +
N2, G + N1 + N2 a kontrola.
Obecně zhodnoceno, odrůda Passandra vykázala vyšší výšku rostliny, počet listů,
obsah čerstvé a suché hmoty v listech. Na druhé straně odrůda Girola dosáhla ve 2. roce
průkazně vyšší rozsah listové plochy. Ranost a celkový výnos byl u obou odrůd srovnatelný.
Tyto výsledky ukazují vliv genotypu na růstové parametry. Přitom Girola dosáhla průkazně
vyšší délky plodů, jejich průměru a hmotnosti. Passandra ovšem vykazovala v obou sezónách
průkazně vyšší počet plodů na rostlinu a současně vyšší zastoupení jakosti „Výběr“ v roce
2007.
Dusík, fosfor a hořčík nevykazoval rozdílný obsah v listech obou odrůd. Obsah
draslíku byl průkazně vyšší u odrůdy Girola v roce 2006 a naopak u odrůdy Passandra v roce
2007. Obsah dusíku v plodech odrůdy Passandra a P, K, Ca v plodech odrůdy Girola byl vyšší
v roce 2006. Genotyp neovlivnil vitamin C ani TSS obsah. Obsah sušiny v plodech odrůdy
Passandra byl průkazně vyšší v obou letech; celkový obsah cukrů v roce 2006. Obsah
dusičnanů byl průkazně vyšší v plodech odrůdy Girola, ale stále pod hygienickým limitem.
Lze shrnout, že rozdíly mezi odrůdami korelovaly s rozdíly danými jejich genotypem. Odrůdy
se lišily v produktivitě výnosu a kvalitě.
Vliv rozdílných substrátů přinesl poznatek, že rostliny pěstované v rašelinovém
substrátu dosáhly průkazně vyššího počtu listů na rostlinu a obsahu sušiny v listech. Naopak
listová plocha i čerstvá hmotnost listů byla mezi sezónami variabilní. Mírný průkazný nárůst
byl zjištěn v případě ranosti rostlin pěstovaných v čedičové plsti. Oproti tomu byl zjištěn
průkazně vyšší výnos – v roce 2006 v rašelinovém a v roce 2007 v čedičovém substrátu.
Rostliny v čedičové plsti dosáhly průkazně vyšší délky a hmotnosti. Ovšem okurky
z rašelinového substrátu dosahovaly průkazně vyšší počet plodů na rostlinu a zastoupení
„Výběru“ v roce 2006, na rozdíl od výsledku v další sezóně. Obsah minerálních látek,
vitaminu C, cukrů i TSS v plodech nebyl mezi substráty rozdílný, s několika výjimkami.
Plody rostlin z čedičové plsti dosáhly průkazně vyšší obsah sušiny a NO3- v roce 2006.
Z tohoto hlediska lze shrnout, že rozdíl v látkovém složení plodů závisí jen zčásti na použitém
pěstebním substrátu.
Rostliny inokulované N-Fixujícími bakteriemi nebo houbou Glomus nevykazovaly
rozdíly v růstu, ranosti, délce a průměru plodů, počtu plodů na rostlinu v obou letech,
s několika výjimkami, oproti neošetřené kontrole. Na druhé straně inokulace pozitivně
přispěla k zlepšení růstových parametrů a výnosu, i když výsledky nebyly signifikantní. Je
třeba poukázat na fakt, že mikroorganismy byly vystaveny v hydroponickém systému řadě
nepříznivých faktorů, jako kyselé pH, dostatečný obsah živin v živném roztoku, což mohlo
vést ke snížení efektu inokulace.
Dále také, všechna inokulovaná ošetření vedla k vyššímu, i když neprůkaznému zvýšení
výnosu. Toto navýšení dosáhlo 8 a 21 % v 1., respektive 2. sezóně. Také bylo doloženo
signifikantní zvýšení průměrné hmotnosti plodů rostlin inokulovaných samostatným
Azotobacter nebo v kombinaci s Azospirillium, ve srovnání s kontrolou v roce 2006.
Inokulace rostlin vedla k zvýšení tržního výnosu, obsahu makroprvků v listech a plodech
obou odrůd, s několika výjimkami.
Obsah vitaminu C v plodech byl průkazně nejvyšší při inokulaci Glomus + Azospirillum.
Také obsah celkových rozpustných solí v plodech byl vyšší při inokulaci Glomus +
Azotobacter + Azospirillum v roce 2006. Na druhé straně průkazné snížení sušiny plodů bylo
zjištěno ve variantě Azotobacter a Azospirillum v roce 2007, ve srovnání s kontrolou. Obsah
nitrátů byl průkazně zvýšen při inokulaci Azotobacter nebo Azospirillum oproti variantě
Glomus nebo kontrole v roce 2006.
Interakce mezi odrůdou, substrátem a inokulací ukázaly řadu kladných vlivů na většinu
růstových charakteristik rostlin okurek. Kupříkladu N-Fixers samostatně nebo v kombinaci s
Glomus zvýšily růst rostlin v rašelinovém substrátu. Okurky v rašelině inokulované Glomus +
Azotobacter nebo Azospirillum měly průkazně vyšší celkový výnos v roce 2006 oproti
kontrole. Lze shrnout, že výnos a růst odrůdy Passandra v rašelinovém substrátu byl pozitivně
ovlivněn inokulací mikroorganismy.
Listy u obou odrůd dosáhly průkazně vyššího obsahu N, K, Ca a Mg při pěstování
v rašelině v roce 2007. Ani odrůda ani substrát jako samostatný faktor neměly vliv na obsah
vitaminu C nebo TSS, mimo několika výjimek. Obsah sušiny, nitrátů a celkových cukrů
v plodech odrůd nebyl rozdílný mezi oběma substráty, mimo několika případů v roce 2007.
Ovšem inokulace s Glomus + Azospirillum zvýšila obsah vitaminu C v plodech obou odrůd,
především v rašelinovém substrátu. Podobně, TSS obsah byl průkazně vyšší v plodech rostlin
z rašeliny, pokud byly rostliny inokulovány Glomus + Azotobacter + Azospirillum. Z tohoto
hlediska lze konstatovat, že inokulace zlepšila nutriční hodnotu plodů oproti neošetřené
kontrole. Toto konstatování odpovídá faktu, že symbiotické mikroorganismy mají podpůrnou
roli v rámci rostlinného metabolismu v listech a výhonech inokulovaných rostlin.
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VII. APPENDIX
Table 9. Effect of interaction between cultivar x microorganisms inoculation on plant
height (m) of cucumbers in 2006 and 2007
Table 10. Effect of interaction between substrate x microorganisms inoculation on plant
height (m) of cucumbers in 2006 and 2007
Table 11. Effect of interaction between cultivar x microorganisms inoculation on leaf
area (cm2) of cucumbers in 2006 and 2007
Table 12. Effect of interaction between substrate x microorganisms inoculation on leaf
area (cm2) of cucumbers in 2006 and 2007
Table 13. Effect of interaction between cultivar x microorganisms inoculation on number
of leaves of cucumbers in 2006 and 2007
Table 14. Effect of interaction between substrate x microorganisms inoculation on
number of leaves of cucumbers in 2006 and 2007
Table 15. Effect of interaction between cultivar x microorganisms inoculation on fresh
weight (g/leaf) of cucumbers in 2006 and 2007
Table 16. Effect of interaction between substrate x microorganisms inoculation on fresh
weight (g/leaf) of cucumbers in 2006 and 2007
Table 17. Effect of interaction between cultivar x microorganisms inoculation on dry
weight (g/leaf) of cucumbers in 2006 and 2007
Table 18. Effect of interaction between substrate x microorganisms inoculation on dry
weight (g/leaf) of cucumbers in 2006 and 2007
Table 21. Effect of interaction between cultivar x microorganisms inoculation on early
yield (kg.m-2) of cucumbers in 2006 and 2007
Table 22. Effect of interaction between substrate x microorganisms inoculation on early
yield (kg.m-2) of cucumbers in 2006 and 2007
Table 23. Effect of interaction between cultivar x microorganisms inoculation on total
yield (kg.m-2) of cucumbers in 2006 and 2007
Table 24. Effect of interaction between substrate x microorganisms inoculation on total
yield (kg.m-2) of cucumbers in 2006 and 2007
Table 27. Effect of interactions between cultivar x microorganisms inoculation on fruit
length (mm) of cucumbers in 2006 and 2007
Table 28. Effect of interactions between substrate x microorganisms inoculation on fruit
length (mm) of cucumbers in 2006 and 2007
Table 29. Effect of interactions between cultivar x microorganisms inoculation on fruit
diameter (mm) of cucumbers in 2006 and 2007
Table 30. Effect of interactions between substrate x microorganisms inoculation on fruit
diameter (mm) of cucumbers in 2006 and 2007
Table 31. Effect of interactions between cultivar x microorganisms inoculation on fruit
number per plant of cucumbers in 2006 and 2007
Table 32. Effect of interactions between substrate x microorganisms inoculation on fruit
number per plant of cucumbers in 2006 and 2007
Table 33. Effect of interactions between cultivar x microorganisms inoculation on fruit
weight (g) of cucumbers in 2006 and 2007
Table 34. Effect of interactions between substrate x microorganisms inoculation on fruit
weight (g) of cucumbers in 2006 and 2007
Table 37. Effect of interaction between cultivar x microorganisms inoculation on fruit
yield of grade "Extra" (kg.m-2) in 2006 and 2007
Table 38. Effect of interaction between substrate x microorganisms inoculation on fruit
yield of grade "Extra" (kg.m-2) in 2006 and 2007
Table 39. Effect of interaction between cultivar x microorganisms inoculation on fruit
yield of "Class A" (kg.m-2) in 2006 and 2007
Table 40. Effect of interaction between substrate x microorganisms inoculation on fruit
yield of "Class A" (kg.m-2) 2006 and 2007
Table 41. Effect of interaction between cultivar x microorganisms inoculation on fruit
yield of "Class B" (kg.m-2) in 2006 and 2007
Table 42. Effect of interaction between substrate x microorganisms inoculation on fruit
yield of "Class B" (kg.m-2) in 2006 and 2007
Table 43. Effect of interaction between cultivar x microorganisms inoculation on
unmarketable fruit yield (kg.m-2) in 2006 and 2007
Table 44. Effect of interaction between substrate x microorganisms inoculation on
unmarketable fruit yield (kg.m-2) in 2006 and 2007
Table 47. Effect of interaction between cultivar x microorganisms inoculation on
nitrogen content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 48. Effect of interaction between substrate x microorganisms inoculation on
nitrogen content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 49. Effect of interaction between cultivar x microorganisms inoculation on
phosphorus content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 50. Effect of interaction between substrate x microorganisms inoculation on
phosphorus content (g/100g dry matter) of cucumbers leaves in 2006 and 2007
Table 51. Effect of interaction between cultivar x microorganisms inoculation on
potassium content (mg.kg-1 fresh weight) of cucumbers leaves in 2006 and
2007
Table 52. Effect of interaction between substrate x microorganisms inoculation on
potassium content (mg.kg-1 fresh weight) of cucumbers leaves in 2006 and
2007
Table 53. Effect of interaction between cultivar x microorganisms inoculation on calcium
content (mg.kg-1 fresh weight) of cucumbers leaves in 2006 and 2007
Table 54. Effect of interaction between substrate x microorganisms inoculation on
calcium content (mg.kg-1 fresh weight) of cucumbers leaves in 2006 and 2007
Table 55. Effect of interaction between cultivar x microorganisms inoculation on
magnesium content (mg.kg-1 fresh weight) of cucumbers leaves in 2006 and
2007¨
Table 56. Effect of interaction between substrate x microorganisms inoculation on
magnesium content (mg.kg-1 fresh weight) of cucumbers leaves in 2006 and
2007
Table 59. Effect of interaction between cultivar x microorganisms inoculation on
nitrogen content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 60. Effect of interaction between substrate x microorganisms inoculation on
nitrogen content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 61. Effect of interaction between cultivar x microorganisms inoculation on
phosphor content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 62. Effect of interaction between substrate x microorganisms inoculation on
phosphor content (g/100g dry matter) of cucumber fruits in 2006 and 2007
Table 63. Effect of interaction between cultivar x microorganisms inoculation on
potassium content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and 2007
Table 64. Effect of interaction between substrate x microorganisms inoculation on
potassium content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and 2007
Table 65. Effect of interaction between cultivar x microorganisms inoculation on calcium
content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and 2007
Table 66. Effect of interaction between substrate x microorganisms inoculation on
calcium content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and 2007
Table 67. Effect of interaction between cultivar x microorganisms inoculation on
magnesium content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and
2007
Table 68. Effect of interaction between substrate x microorganisms inoculation on
magnesium content (mg.kg-1 fresh weight) of cucumber fruits in 2006 and 2007
Table 71. Effect of interaction between cultivar x microorganisms inoculation on vitamin
C (mg.kg-1) of cucumber fruits in 2006 and 2007
Table 72. Effect of interaction between substrate x microorganisms inoculation on
vitamin C (mg.kg-1) of cucumber fruits in 2006 and 2007
Table 73. Effect of interaction between cultivar x microorganisms inoculation on TSS
(%) of cucumber fruits in 2006 and 2007
Table 74. Effect of interaction between substrate x microorganisms inoculation on TSS
(%) of cucumber fruits in 2006 and 2007
Table 75. Effect of interaction between cultivar x microorganisms inoculation on dry
matter (%) of cucumber fruits in 2006 and 2007
Table 76. Effect of interaction between substrate x microorganisms inoculation on dry
matter (%) of cucumber fruits in 2006 and 2007
Table 77. Effect of interaction between cultivar x microorganisms inoculation on NO3-
(mg.kg-1) of cucumber fruits in 2006 and 2007
Table 78. Effect of interaction between substrate x microorganisms inoculation on NO3-
(mg.kg-1) of cucumber fruits in 2006 and 2007
Table 79. Effect of interaction between cultivar x microorganisms inoculation on total
sugars (mg.kg-1) of cucumber fruits in 2006 and 2007
Table 80. Effect of interaction between substrate x microorganisms inoculation on total
sugars (mg.kg-1) of cucumber fruits in 2006 and 2007

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