Egypt. J. Agric.

Res, 86(3), 2008

1197

PHOTOSYNTHESIS, LEAF AREA INDEX AND PRODUCTIVITY OF
TOMATO (Lycopersicon esculentum Mill.) GERMPLASM

Milton E. Pereira Flores
1
; Derly J. Henriques da Silva
1
; Marco Oliva
2
; Francisco
X.R. Do Vale
3
; Jorge G. Aguilera
4
; Ahmed Y. Elsayed
4,5



1
Plant Sciences,
2
Plant Physiology,
3
Phytopathology and
4
Genetics and
Breeding Depts., University Federal of Viosa, MG, Brazil.
5
Hort. Res. Inst., Agric. Res. Center, Giza, Egypt.

(Author for corresponding: miltonpereira2001@yahoo.com)

ABSTRACT

Nine accessions of tomato (Lycopersicon esculentum Mill.)
indeterminate growth habit belonging to BGH-UFV bank with high
variability were evaluated for leaf net photosynthesis (A), leaf area
index (LAI), total dry matter production, dry matter partitioning and
productivity. Santa Clara cultivar was used as well-adapted standard
tomato variety with high production, LAI and total dry matter. The
genotypes were grown under optimal irrigation and nutrition
conditions. Optimal environmental conditions occurred over all
experiment. The results showed significant genetic variability for all
the evaluated traits. No correlation was found between A, total dry
matter and productivity. Positive correlation between total dry
matter, production and harvest index, but A correlated negatively
with LAI. Favorable combination of physiological traits with
production was found in BGH-320 and BGH-1020 accessions.

INTRODUCTION

The physiological breeding has considered the study of morpho-physiologic
characteristic with the aim of identifying quantitative traits or combinations used in
indirect selection with high efficiency (3, 4). In this context and in different cultivars has
been frequent the seeking for genetic variability of leaf net photosynthesis (A) measured
by CO
2
gas exchange and its effective contribution with improved the production (1, 15,
17, 21, 22, 28). Early research in this area has related linear increases, with different
rates, from A, harvest index (HI), stomatal conductance and chlorophylls content in
genetic improvement of cultivars as soybean (22), rice and wheat (21). In contrast, other
studies (28, 30) have indicated that the large increases in production do not require
increase of A, but only of the morpho-agronomic characteristics as biomass, plant
architecture and size, colour of the canopy and the harvest index, resulting in productive
cultivars with high leaf area index (LAI) and reduced photosynthetic capacity per leaf unit
(9, 28). It has been reported genetic variability, heritability and transgressive F1 hybrids
in A for diverse environmental conditions in tomato (5, 6, 11, 31). However, the
correlations between A and yield, total dry mass, content of chlorophyll and fruit quality
Egypt. J. Agric. Res, 86(3), 2008

1198

are contradictory among scientific research and have needing for more knowledge in this
area.

Thus, the result of the present paper arise the knowledge of the genetic variability of
photosynthetic capacity and its relation to production, total dry matter, IAF and harvest
index of tomato plants.


MATERIALS AND METHODS

In this study it was used 9 tomato accessions (Lycopersicon esculentum Mill.) and
Santa Clara cultivar belonging to Vegetables Germplasm Bank of University Federal of
Viosa, BGH-UFV (http://www.ufv.br/bgh/files/pag/tomate.htm) with high morpho-
agronomic variability. 'Santa Clara' was used as a standard well-adapted and productivity
reference of the Santa Cruz type cultivars and because of its wide diffusion in Brazil (26,
29). The experiment was conducted in greenhouse under favourable environmental,
nutrient and irrigation conditions (8, 12, 14). The soil was plough up with 8 kg MS
-2
m of
manure "tanned" until 0.40 m depth, to improve the porosity of soil for avoid any
hypoxic conditions than affect severely the photosynthesis along the A assessment. The
irrigation was calculated using lysimeter (18) and the average water soil potential in 10
and 30 cm of depths was 10.63 8.34 and 27.8 48.4 Kpa, respectively (19).

The seedlings were transplanted at 31 days-old (four to five extended leaves), 0.5 m
x 1.0 m space, a randomized block design with five replications and 2 plants per block
was used. The plants were grown in vertical system with one shoot per plant and pruned
above the second pair of leaves of the fifth bunch. The A was measured with IRGA
analyzer (LI-6400, LI-COR Biosciences, Inc. Lincoln, Nebraska) in the second pair of
leaflets of the fourth leaf from 7:30 to 11:00 AM at 51 to 56 day after transplanting. The
IRGA was configured as following: air flow of 500 mmol s
-1
, 1200 mol photons m
-2
s
-1

and 25
o
C for the chamber and leaves temperatures (15, 16, 24). This measurement
period was chosen in agreement with (11) that found in 54 DAT the highest A values in
12 tomato genotypes, measurements over different development stages. The fourth leaf
was used in previous test which had highest and stable value of A. The leaf area index
(LAI), leaf area (LA), total dry matter (TDM) and harvest index (HI) were calculated by
conventional methods and expresses in standard units. The root biomass was extracted
of a soil cube (30x30x30 cm) by washing with running water on sieve.

Statistical Analysis: The data was submitted to analyzed of variance using F-test (p
<0.01), discrimination of means by Tukey (p <0.05) and correlations by Pearson with
SAEG 0.9 statistical software.







Egypt. J. Agric. Res, 86(3), 2008

1199

RESULTS AND DISCUSSION


Production
The production, as it predicted, had significant variation among the genotypes (F
test, p <0.01). The BGH-1020, `Santa Clara and BGH-994 were superiors than other
genotypes. Contrary, the BGH-322 and BGH-406 had lowest production (two kg pta
-1
)
representing 31% or less than productive genotypes. The productivity of Santa Clara,
BGH-1020, BGH-994, BGH-970 and BGH-320 was equivalent to 116; 117; 105; 84 and 84
ton ha
-1
, respectively. This production was superior to the average tomato yield in Brazil
60 ton ha
-1
(20) thus the best accessions could be used as progenitors to increase
tomato productivity.

Leaf net photosynthesis
The leaf net photosynthesis (A) of the genotypes was wide variability (F test, p
<0.01) between 25 and 17 mol CO2 m
-2
s
-1
(Table 1); these values was related in early
research for accessions (11) and hybrids (31) of the specie. The accessions BGH-320
and BGH-1020 had largest A, with 25 and 23 mol CO
2
m
-2
s
-1
, respectively, although
BHG-1020 did not differ from other accession. All the accessions were higher than the
commercial tomato `Santa Clara which reached 17 mol CO
2
m
-2
s
-1
.

Table 1, Santa Clara had lowest A value whereas BGH-320 with the highest A value it
means less production than Santa Clara or BGH-994. The Pearson correlation (Table 2),
did detect any relation between these traits, indicating there is no direct influence of
photosynthesis on the production in tomato. This results is coherent with previous
research with tomato (5) but is contrary with other pre-breeding research of tomato
germplasm (11) which related positive correlation between A and total dry mass and
production. These antagonist reports may be understand due to A is highly variable over
development stages of plant not integrative measurement of whole net photosynthesis,
and highly influenced by the environmental conditions, in addition, the true maximum
values of A is ever recorded and not standard homogenized protocols are applied for the
A assessment. In this experiment, it was applied similar development stage (11) but not
similar experimental conditions.

On the other hand, the inheritance of this trait have related transgressive
inheritance, significant general combining ability and the potential selection of A in
tomato (31, 5) which suggest this character is controlled by additive gene action. This
fact implicates the possibility of accessions BGH and particularly BGH-320 and BGH-1020
can be used as parents to increase A in tomato breeding.

Total dry matter and Harvest Index
The total dry matter (TDM) and harvest index (HI) had wide variation among the
genotypes (F test, p <0.01). The total dry matter of BGH-1020 was superior to all the
other genotypes, except BGH-994 and Santa Clara. These genotypes had superior
production too. Correspondingly the BGH-406 had the less TDM and production. The
others accessions showed moderate TDM and production (Table 1). Indeed the Pearson
correlation was positive for relation of TDM and Production (Table 2). Contrary didnt
Egypt. J. Agric. Res, 86(3), 2008

1200

have correlation among A, production and total dry matter as reported in previous study
in pre-breeding tomato genotypes (11). Our data coincident with the results obtained
from cotton (17) and tomato (6, 31). In these cultivars didnt found any positive
correlation among these traits.

The harvest index varied significantly (F test, p < 0.01) between 25 and 64% (Table
1). The accessions BGH-320 with HI 64% allocated 10% more than the dry matter fruit
reached for Santa Clara between 10 and 40% of the rest accessions. The value of HI for
tomato plant ranged between 60 and 70% (12) despite of the experiment conditions
were similar.

The high TDM and its correlated with the production implies plants with high TDM,
potentially may allocate more units of mass to fruit, even though the HI being the same
if compared with other plant that has less TDM. The BGH-1020, Santa Clara and BGH-
994 with the largest TDM had the highest productions too, but its HI was 10 to 16%
units below BGH-320 had IH 64% and 6.5% lower production. In this case, the
production of BGH-320 would be limited by its TDM. Similarly, the BGH-322 and BGH-
406 with the lower production had TDM and HI of 25 and 40% which means that high
TDM improved the probability to obtain high HI in tomato breeding as well as it related in
other cultivars (14, 28, 30).

On the other hand, the lack of correlation between A and productive traits, suggest
that the A variability will be important only if exist inheritance and significant correlation
with production or other traits. Between the F1 hybrids, consequently the A variability
may be desirable over the correlation with production, for the improvement the
productive efficiency of the tomato plants.

Leaf area Index (LAI)
There was significant variation of LAI among genotypes (F test, p <0.01) at 83 DAT
and the EPC (Table 2). `Santa Clara, with the least A had a LAI superior to all accessions
at 83 DAT and the EPC, but only has different with BGH-320, BGH-987, BGH-970 and
BGH- 406 accessions at the EPC (Table 1).

This superiority of the Santa Clara in LAI over the accessions is consistent with the
fact of the selection for production in the new cultivars would reduce A and increase IAF
to compensate the reduced exchange of CO
2
per unit area (28,30) as effectively was
observed in Santa Clara (Table 1).

The BGH-320 with high A had the lowest LAI to 83 DAT (1.4) and intermediate value
in the EPC (2.3), but was significantly lower than the IAF in Santa Clara. BGH-1020 with
the largest production had intermediate values of IAF in both evaluations, but only
reached the EPC values equal to `Santa Clara.

In this context the ideal limit of LAI in tomato is between 3.0 and 3.5 (12) even
though in most commercial cultivars ranges between 90 and 120 DAT, from which occurs
reduction of leaf area by the end of the production cycle (7, 10, 27). Above this optimal
level of LAI, the shelf-shaded decrease the efficiency in the use of radiation, this effect
Egypt. J. Agric. Res, 86(3), 2008

1201

decrease the average rate of irradiance consequently more leaves below the light
compensation point, increase the maintenance respiration, leaf senescence and the risk
to the development of diseases (2, 10, 12, 23).

The continuing increase of LAI in accessions, contrary to that observed in Santa
Clara, indicates that there was no involvement of any major fraction of leaf area early, in
fact many accession had active bottom leaves in end harvest. The elimination of bottom
leaves is recommended practices to maintain the source/sink relation, however this
practices is expensive physiologically and economically. This cost is associated with
exuberant cultivars like Santa Clara, even that could be unnecessary if the plant would
combine high A values per surface leaf unit with high HI like BGH-320.

This expressive inverse association between the IAF and A, observed in `Santa Clara
and BGH-320 indicate the importance of considering together the IAF, A and HI in the
development of news productive cultivars. This inverse association between the IAF and
A, was early reported in cereals, in which were associated small leaves with
photosynthetic activity maximized per leaf unit (30).


CONCLUSION

There was genetic variability for all the physiological traits assessment in the accessions
of BGH-UFV. The leaf net photosynthesis (A) had negative correlation with desirable
agronomic and physiological characteristics like production and leaf area index (LAI). The
favourable combination of A, IAF and HI found in BGH-320, which can be used as
promises genotype for breeding research aims development a new tomato ideotype, by
combining highest A, HI with lower LAI.

REFERENCES

1. Agnihotri, R.K., Chandra, S., Sharma, S., And Palni, L.M.S., G.B. Genetic
variability in photosynthesis and chlorophyll content of various landraces of
upland rice Plant Institute of Himalayan Environment and Development, Kosi-
Katarmal, Almora 263643, ndia. IRRN 25.2. August 2000. 2p.
2. Andriolo, J.L. Fisiologia das culturas protegidas. Santa Maria: ed. da UFSM,
1999. 142p.
3. Araus, J.L; Casadesus, J.B.; Nachit, M.M.; Villegas, D.; Aparicio, N. And Royo, C.
Some remarks on ecophysiological traits for breeding. Noureaux defies
Zaragoza; CIHEAM-IAMZ. 2000. 620 p.
4. Borem, A; Miranda, G.V. Melhoramento de plantas. 4. ed.Viosa:UFV, 2005. 525
p.
5. Dijk, S.J.D.V. Inheritance of net photosynthesis, dark respiration, stomatl
resistance and related characters in tomato (Lycopersicon esculentum Mill.)
under low energy conditions Euphytica. Volume 36, Number 1/ January, 1987.
709-716 p.
Egypt. J. Agric. Res, 86(3), 2008

1202

6. Dijk, S.J.D.V.; Maris, J.A. Differences between genotypes in net photosynthesis
and dark respiration under low light intensity and low night temperatures.
Journal Euphytica. Volume 34, Number 3/ November, 1985. 709-716 p.
7. Dogliotti, S. Bases fisiolgicas del crecimiento y desarrollo del cultivo del tomate
(Lycopersicon esculentum Mill.). Fisiologa de cultivos. Universidad de la
Repblica Facultad de Agronomia. 2004. 18 p.
8. Fontes, P.C.R. Sugestes para adubao para hortalias. Recomendaes para o
uso de corretivos e fertilizantes em Minas Gerais. 5 aproximao. p. 171-174.
Viosa. 1999. 300 p.
9. Geber, A.M.; Griffen, R.L. Inheritance and natural selection on functional traits.
Int. J. Plant Sci. 164(3 Suppl.):S21S42. 2003. University of Chicago.
10. Guerra, H.M.; Borrego, E.F.; Oyervides, G.A.; Fernandez, B.J.M. Evaluacin de
gentipos de tomate considerando critrios fisiolgicos, fenolgicos y de
rendimiento, bajo condioes de alta temperatura en invernadero. AGRARIA
UAAAN VOL 15 NUM. 1 DE 1999. UNIVERSIDAD AUTONOMA AGRARIA ANTONIO
NARRRO Buenavista, Saltillo, Coah., Mxico. 55 77 p.
11. Guimares, M.D.A. Influencia da poda apical e da posio do cacho de tomateiro
no crescimento da planta e na qualidade de frutos. Viosa: UFV. 2004. 93p.
(Tese de Mestrado).
12. Heuvelink, E. Dry matter partitioning in tomato plant: one common assimilate
pool?. J. Experimental Botany, 46, p. 1025-1033, 1995.
13. Heuvelink, E. Tomatoes. CABI publishing. 2005. 339p.
14. Heuvelink, E; And Buiskool, R.P.M. Influence of sink-source interaction on dry
matter production in tomato. Annals of Botany 75: 381-389, 1995.
15. Hunt, S. Measurements of photosynthesis and respiration in plants. Physiology
plantarum 117: 314-325. 2003.
16. Licor. Using the LI-6400. Portable Photosynthesis System Manual. Version 5.
2004, LI-COR, Inc. 1106 p.
17. Lu, Z.; Percy, G.R.; Qualset, O. C.; Zeiger, E. Stomatal conductance predicts
yields in irrigated Pima cotton and bread wheat grow at high temperatures.
Journal of Experimental Botany, Vol. 49, Special Issue, pp. 453 460, March
1998.
18. Machado E.R.; Matos A. Construo e instalao de um lismetro com sistema de
Drenagem. Revista Brasileira de Agrometeorologia, Santa Maria, v. 9, n. 1, p.
147-151, 2001
19. Marouelli, W. A., Silva, W. L. C., Silva, H. R. Manejo de irrigao em hortalias.
5.ed. Braslia: Empresa Brasileira de Pesquisa Agropecuria - CNPH, 1996. 71p.
20. Mattedi, A.P.; Soares, B.O.; Almeida, V.S., Grigolli, J.F.J.; Silva, L.J.; Silva, D. J.
H. Introduo cultura do tomateiro. In: Silva, Derly Jos Henriques; Vale,
Francisco Xavier Ribeiro. (Org.). Tomate - Tecnologia de produo. Visconde do
Rio Branco: Suprema, 2007, v. , p. 85-99.
21. Ming, Z.; Lafitte, R.; Sacks, E.; Dimayuga, G.; Zhao, X. Variation in leaf
photosynthesis among wild species in Genus Oryza and among the progeny of
selected crosses of the wild species with a Riece cultivar. International Rice
Research Institute. 2005. 5p.
Egypt. J. Agric. Res, 86(3), 2008

1203

22. Morrison, M.J.; Voldeng, H.D.; Cober, E.R. Physiological changes from 58 years
of genetic improvement of short-season Soybean cultivars in Canada. Agronomy
Journal, Vol. 91: 685-689. July-August 1999.
23. Oliveira, C.A.; Maroulli, W.A.; Santos, J.R.M; Boiteux, L.S. Produao de
esclerodios de Sclerotinia sclerotiorum e severidade de oidio em cultivares de
ervilha sob diferentes laminas de irrigao. Horticultura Brasileira, Braslia, v. 18,
n.1, p. 16-20, maro 1999.
24. Pachepsky, L.B; Acock, B. A model 2DLEAF of gas exchange: development,
validation an ecological application. Ecological Modelling 93 (1996) 1-18.
25. Paz, L.G., Matos, M.M.V.L. 1985. A fotossntese e a produtividade das
forrageiras. In: UNIVERSIDADE FEDERAL RURAL DE PERNAMBUCO, 1985,
Recife. Anais... Recife, 1985, v.8, n.10, p.103-20.
26. Peixoto, J.R.; Oliveira, C.M.D; Silva, R.P.D. Angelis, B.D.; Ceclio Filho, A.B.
Avaliao de gentipos de tomateiro tipo Santa Cruz no perodo de inverno em
Araguari, MG. Pesquisa Agropecuria Brasileira, Braslia, v.34, n.12, p.2247-2251,
dezembro. 1999
27. Peluzio, J. M. Crescimento e partiao de assimilados em tomateiro (Lycopersicon
esculentum Mill.) aps a poda apical. Viosa: UFV, 1992. 49p. (Tese de
Mestrado).
28. Richards, R. A. Selectable traits to increase crop photosynthesis and yield of
grain crops. Journal of Experimental Botany, Vol. 51. GMP Special Issue. Pp.
447-458, February 2000.
29. Santos, M.M.D.; Noronha, J.F.D. Diagnstico da cultura do tomate de mesa no
Municpio de Goianpolis, Estado de Gois, Brasil. Pesquisa Agropecuria
Tropical, 31(1): 29-34, 2001 31.
30. Sharma, N.P.; And Ghildiyal, C. M. Potential targets for improvement
photosynthesis and crops yield. CURRENTE SCIENCE, vol.88, no. 12, 25 june
2005. 1918-1925 p.
31. Weerasinghe, O.R.; Perera, A.L.T.; De Costa, W.A.J.M.; Jinadase, D.M.;
Vishnukanthasingham, R. Production of tomato hybrids for dry zone conditions
of Sri Lanka using combining ability analysis, heterosis and DNA testing
procedures. Tropical Agricultural Research Vol. 16: 79-90 (2004).














Egypt. J. Agric. Res, 86(3), 2008

1204

,
(Lycopersicon esculentum Mill.)


1

1

2

3

4


45
.


1

2

3

4
- - -
.
5
.


(Lycopersicon esculentum Mill.)
) A ( ) LAI (
.

.
. .

A LAI . A

. BGH-320 BGH-1020
.





















Egypt. J. Agric. Res, 86(3), 2008

1205

Table 1: Agronomic and physiological traits of the BGH accessions and Santa Clara`
grown in greenhouse. Viosa-MG, Brazil 2006.

Accession
or
cultivar
Production
kg pta
-1

A*
Total Dry
Matter
g pta
-1

Harvest
Index
%
Leaf Area Index
m2 m
-2

83 DAT* EPC*
BGH-320
BGH-1020
BGH-987
BGH-322
BGH-970
BGH-406
BGH-349
BGH-975
BGH-994
4.20 bc
5.84 a
3.80 d
1.81 e
3.96 d
1.82 e
3.40 d
4.20 cd
5.24 ab
24,7 a
22.6 ab
22.0 b
21.7 b
21.3 b
21.0 b
21.0 b
20.7 b
20.4 b
359 e
590 a
373 e
395 de
463 bcd
263 f
316 ef
459 cd
547 ab
64 a
49 bc
50 bc
25 e
46 cd
40 d
55 b
50 bc
48 bc
1.4 d
1.8 bcd
1.7 cd
1.6 cd
1.6 cd
1.6 cd
1.7 cd
2.2 bcd
2.3 b
2.3 cd
3.2 ab
2.3 cd
3.0 ab
2.8 bc
1.9 d
3.3 ab
3.2 ab
3.2 ab
Santa Clara 5.80 a 17.2 c 514 abc 54 b 3.8 a 3.6 a
Average 4.07 21.3 427.8 48.1 2.0 2.9
Dms 8.5 5.3 9,4 7.0 16.8 11.8
* A= Leaf Net Photosynthesis (mol CO
2
m
-2
s
-1
). DAT=Day after transplanting and EPC=End
productive cycle. Averages with the same letter on the columns are similar statistically by Tukey
Test (p < 0.05).

Table 2: Pearson correlation (r) for agronomic and physiological traits of BGH
accessions and Santa Clara` grown in greenhouse. Viosa-MG, Brazil
2006.
Tomato
Traits
A

mol CO
2
m
-2
s
-1

Production
g pta
-1

Total Dry
Matter
g pta
-1

Harvest
Index
%
A

Production 0.1170
Total Dry Matter 0.2080 0.7490**
Harvest Index 0.1018 0.6648** 0.0424
LAI 83 DAT - 0.565** 0.3986** 0.3560** 0.0926
LAI EPC - 0.380** 0.4180** 0.2670** 0.0584
LA 83 DAT - 0.5651** 0.3986** 0.3558** 0.0926
LA EPC - 0.4055** 0.4177** 0.6666** -0.0584
** Pearson coefficient significative by t-test (p < 0.01).

A= Leaf Net Photosynthesis. LAI=Leaf

area index. DAT=Days after transplanting. EPC=End productive cycle. LA=Leaf area.