First results on evaluation of Arundo donax L. clones collected in Southern Italy

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Industrial Crops and Products 23 (2006) 212–222 First results on evaluation of Arundo donax L. clones collected in Southern Italy Salvatore Luciano Cosentino , Venera Copani, Giuseppina Marina D’Agosta, Emanuele Sanzone, Mariadaniela Mantineo Sezione di Scienze Agronomiche, Dipartimento di Scienze Agronomiche, Agrochimiche e delle Produzioni animali dell’Universit` a di Catania, via Valdisavoia 5, 95123 Catania, Italy Received 28 March 2003; accepted 22 June 2005 Abstract The research of alternative crops for biomass production for energy indicates giant reed (Arundo donax L.), widespread spontaneous plant in Mediterranean regions, among the species at high aptitude for accumulation of biomass. Within the activity of an E.U. programme (CEE FAIR CT 97-2028 “Giant reed (A. donax L.) Network. Improvement, productivity and biomass quality”, germplasm of giant reed were collected to evaluate potential production and the phenotypic and genotypic variability, the heritability in order to selecting the best genotypes. In 1997 and 1998, trials were carried out in Primosole site (Piana of Catania, sea level, 37 25 N latitude; 15 30 E longitude), utilizing rhizomes of 39 clones collected in Sicily and Calabria. The rhizomes were transplanted in springtime. Phenological (date of flowering), biometrical (stem density, stem height, number of nodes per stem, diameter and thickness of stems, weight of fresh and dry biomass of leaves, stems and inflorescence) and productive (yield) data were measured. Harvest were carried out in February 1988 and 1989. Yield of 39 clones studied was, in the average, 10.6 t ha 1 of dry matter in the first year and 22.1 t ha 1 in the second one. The clone no. 4 (Piazza Armerina) and the clone no. 20 (Capo d’Orlando) maintained their high productive aptitude in both years; they yielded respectively, 13.1 and 14.1 t ha 1 in the first year and 34.2 and 26.9 t ha 1 in the second one. The yield results positively correlated to stem density, stem weight and plant height. Four characters: biomass yield, stem weight, stem density and stem height showed a significant variance among clones without significant interaction with year. Among the eleven characters measured only yield, stem weight, stem density and stem height had moderate heritability (h 2 ), comprised between 23 and 48% showing promise for genetic improvement. © 2005 Elsevier B.V. All rights reserved. Keywords: Arundo donax L.; Clonal species; Genetic diversity; Heritability; Mediterranean environment Corresponding author. Fax: +39 095 234449. E-mail address: [email protected] (S.L. Cosentino). 0926-6690/$ – see front matter © 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.indcrop.2005.06.004

Transcript of First results on evaluation of Arundo donax L. clones collected in Southern Italy

Industrial Crops and Products 23 (2006) 212–222

First results on evaluation ofArundo donax L. clones collectedin Southern Italy

Salvatore Luciano Cosentino∗, Venera Copani, Giuseppina Marina D’Agosta,Emanuele Sanzone, Mariadaniela Mantineo

Sezione di Scienze Agronomiche, Dipartimento di Scienze Agronomiche, Agrochimiche e delle Produzioni animalidell’Universita di Catania, via Valdisavoia 5, 95123 Catania, Italy

Received 28 March 2003; accepted 22 June 2005

Abstract

The research of alternative crops for biomass production for energy indicates giant reed (Arundo donax L.), widespreadspontaneous plant in Mediterranean regions, among the species at high aptitude for accumulation of biomass. Within the activityof an E.U. programme (CEE FAIR CT 97-2028 “Giant reed (A. donax L.) Network. Improvement, productivity and biomassquality”, germplasm of giant reed were collected to evaluate potential production and the phenotypic and genotypic variability,the heritability in order to selecting the best genotypes.

In 1997 and 1998, trials were carried out in Primosole site (Piana of Catania, sea level, 37◦25′N latitude; 15◦30′E longitude),ologicals, weightre carried

eh years;

ld, stemith year.bility (

utilizing rhizomes of 39 clones collected in Sicily and Calabria. The rhizomes were transplanted in springtime. Phen(date of flowering), biometrical (stem density, stem height, number of nodes per stem, diameter and thickness of stemof fresh and dry biomass of leaves, stems and inflorescence) and productive (yield) data were measured. Harvest weout in February 1988 and 1989.

Yield of 39 clones studied was, in the average, 10.6 t ha−1 of dry matter in the first year and 22.1 t ha−1 in the second one. Thclone no. 4 (Piazza Armerina) and the clone no. 20 (Capo d’Orlando) maintained their high productive aptitude in botthey yielded respectively, 13.1 and 14.1 t ha−1 in the first year and 34.2 and 26.9 t ha−1 in the second one.

The yield results positively correlated to stem density, stem weight and plant height. Four characters: biomass yieweight, stem density and stem height showed a significant variance among clones without significant interaction wAmong the eleven characters measured only yield, stem weight, stem density and stem height had moderate heritah2),comprised between 23 and 48% showing promise for genetic improvement.© 2005 Elsevier B.V. All rights reserved.

Keywords: Arundo donax L.; Clonal species; Genetic diversity; Heritability; Mediterranean environment

∗ Corresponding author. Fax: +39 095 234449.E-mail address: [email protected] (S.L. Cosentino).

0926-6690/$ – see front matter © 2005 Elsevier B.V. All rights reserved.doi:10.1016/j.indcrop.2005.06.004

S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222 213

1. Introduction

Giant reed (Arundo donax L.) is a perennial, herba-ceous plant, graminaceous family of grasslands andwetlands occurring over a wide range of climatic habi-tats. It has a C3 photosynthetic cycle, but it has highrates of photosynthesis and productivity similar tothose of C4 species (Christou, 2001).

Thousand of years ago, it was cultivated in Asia,South Europe, North Africa and Middle East andit was introduced and diffused widely in North andSouth America and in Australia in the 19th century(Perdue, 1958; Zohary, 1962). It is used generally tosatisfy local necessities, like as plant tutor, basketsand mats, and mainly musical instruments. In Italy,this species was utilized industrially since 1930, whenSnia-Viscosa registered a trademark to obtain cellulosepasta for the production of rayon viscose and paper(Facchini, 1941).

According to some authors Giant reed is native fromEast Asia (Polunin and Huxley, 1987); it is diffusedin all Mediterranean environment, where usuallydoes not set fruit because the pollen results unfruitful(Boose and Holt, 1999); consequently, the speciesis thought to spread primarily asexually by flooddispersal of stem cuttings and rhizome pieces. In thissituation, the natural variability in existing populationsof clones, as it is known, may occur due to spontaneousmutation followed by natural selection as a responseto climatic stresses and to different environment, orb s ofd

oneo astap ,1 ira,2 ap-te 01h ent(

ec-e o theM reado hatw thev d-e me

a study of the genetic diversity among populationsappears to be useful.

From 1997 to 2000, an European project (FAIR3CT96 2028): “Giant reed (A. donax L.) NetworkImprovement, productivity and biomass quality” hasbeen carried out to obtain useful information on the pos-sibility to utilize this species on wide scale in Europe.Within this framework, in the South of Italy researchwere carried out in order to examine the existing phe-notypic and genotypic variability in the extensivelydiffuse populations ofA. donax in this area.

2. Material and methods

From April the 7th till the 29th 1997, in the South-ern Italy, based on exploration along sides and beds oftorrents and rivers, but also in scattered uncultivatedareas, thirty-nineA. donax L. clones were collected(35 in Sicily region and 4 in Calabria region). The sam-pling sites ranged from 39◦05′N latitude to 17◦07′Elongitude (Crotone) and 37◦04′N latitude to 14◦15′Elongitude (Gela); the minimum distance betweenlocations was greater of 10 km (Fig. 1; Table 1). Ateach location, more than 30 rhizome pieces were col-lected, the stand of giant reed sampled was consideredto be one clone (rhizomes collected from a singleplant).

All clones were brought in a typical Xerofluvent soil(Fierotti, 1988) at the experimental field of the Uni-v oleal

leteb as7 67p enp

dedi db foret andf s-p -n erei

ao the

y transferring part of plant through the usual wayiffusion.

Recently, this species has been indicated likef the most promising for energy and cellulose production for the Southern areas of Europe (Lunnan997; Foti and Cosentino, 2001; Anatoly and Pere002) in relation to its traits: perennial grass, easy ad

ation to different environmental conditions (Vecchiett al., 1996; Mardikis et al., 2001; Christou et al., 20),igh production of biomass, low input requiremVecchiet and Jodice, 1996; Cosentino et al., 1999).

In order to cultivate this species, it should be nssary to breed the most productive genotypes tediterranean environment. However, because spccurs almost exclusively asexually, it is likely tithin the environments colonised by the speciesariability useful for the breeding activity may be most. Therefore, before to start a selection program

ersity of Catania, Faculty of Agriculture (Primosrea, 10 m at sea level, 37◦25′N latitude; 15◦30′E

ongitude).The clones were grown in a randomised comp

lock design with two replications. Each plot w.5 m2 (2.5 m× 3 m); the plant density was 2.lants m−2 (0.75 m between rows, 0.50 m betwelants).

The day after collection, the rhizomes were divin little portions with two or three visible well-formeuds and they were transplanted in each plot. Beransplanting, the soil was ploughed, harrowedertilised with P2O5 (100 kg ha−1) as mineral perphohate. After transplant, nitrogen (80 kg ha−1) as ammoium sulphate and irrigation (30 mm of water) w

mmediately applied.During the second year, in early spring, 80 kg h−1

f nitrogen (as ammonium nitrate) was supplied to

214 S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222

Fig. 1. Map of giant reed collection locations in Sicilia and Calabria regions.

crop. The harvest of stems was carried out from Febru-ary the 26th till March the 4th 1998 and from Februarythe 15th till the 26th 1999, in the first and in the secondgrowing period, respectively.

In both years irrigation was supplied during drysummer period on the whole 300 and 150 mm of waterwere distributed, respectively, for the first and the sec-ond year; weed control has been done manually.

2.1. Data collection

During the 2 years, the main meteorological parame-ters (maximum and minimum air temperature, rainfall,global radiation) were measured by means of a datalogger (CR 10—Campbell Scient. Inc. Logan, Utah),with the relevant probes, which was located close tothe experimental field.

During the cropping period the following phenolog-ical stages were recorded:

• emergence of the first buds from the soil;• full flowering when the 50% of the plants showed

the inflorescence completely open.

The harvest of stems was carried out in winter(February), when the stems had the minimum contentof humidity. In central part of each plot (3 m2), thefollowing measurements were carried out: number ofstems; weight of the above ground fresh biomass.

On five stems randomly selected from each plot, thefollowing characters were measured: height of stems(cm) from the base node to the top node without theinflorescence; number of nodes per stem; diameter(mm) and thickness of stems (mm) in three positionsin the middle part of the basal, medium and top intern-odes; weight of fresh biomass (g) of leaves, stems andinflorescence.

A subsample of leaves, stems and inflorescence wasdried at 105◦C until reaching a constant mass, to deter-minate the percentage of dry biomass.

2.2. Statistical procedures

2.2.1. Comparison of clonesAll measured and derived data were subjected to

analysis of variance (ANOVA) separately for each year,according to the experimental layout. In order to com-

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Table 1List of collected clones

Clones Name Geographiccoordinates

Altitude

LatitudeN

LongitudeE

a.s.l.m.

1 S.S. 417 BR 37◦20′ 14◦45′ 2002 S.S. 417

Caltagirone37◦14′ 14◦31′ 608

3 Caltagirone-SantoPietro

37◦07′ 14◦32′ 313

4 PiazzaArmerina-Fegotto

37◦23′ 14◦22′ 697

5 Bivio Ramata 37◦31′ 14◦18′ 6706 Piedimonte etneo 37◦48′ 15◦10′ 3487 Passopisciaro 37◦50′ 15◦08′ 5508 Maniace 37◦53′ 14◦47′ 7879 Bicocca 37◦30′ 15◦04′ 7

10 Fondachello 37◦45′ 15◦11′ 111 Santa Tecla 37◦37′ 15◦10′ 2212 Fontane Bianche 36◦57′ 15◦12′ 1713 Lentini 37◦17′ 15◦00′ 5314 CT-RG bivio

Vittoria36◦57′ 14◦32′ 168

15 Modica 36◦50′ 14◦46′ 29616 Pozzallo 36◦43′ 14◦51′ 2017 Gela 37◦04′ 14◦13′ 4618 Biancavilla 37◦38′ 14◦52′ 51519 Tortorici 38◦ 01′ 14◦49′ 45020 Capo D’Orlando 38◦08′ 14◦43′ 821 S. Stefano di

Camastra38◦00′ 14◦20′ 70

22 Cefalu 38◦01′ 14◦00′ 1623 Roccalumera 37◦58′ 15◦23′ 724 Villafranca 38◦14′ 15◦26′ 2225 Milazzo 38◦13′ 15◦14′ 126 Caltanissetta 37◦29′ 14◦02′ 56827 Agrigento 37◦19′ 13◦35′ 23028 Ribera 37◦29′ 13◦15′ 22329 Menfi 37◦36′ 12◦58′ 11930 Licata 37◦06′ 13◦56′ 831 Trabia 38◦00′ 13◦38′ 5032 Capaci 38◦10′ 13◦14′ 5133 Castellammare 38◦01′ 12◦53′ 2634 Birgi 38◦01′ 12◦32′ 335 Mazara 37◦39′ 12◦35′ 836 Gioia Tauro 38◦25′ 15◦53′ 2937 Lamezia 38◦58′ 16◦18′ 21638 Catanzaro 38◦53′ 16◦65′ 32039 Val di Neto 39◦05′ 17◦07′ 8

pare clones performance, the factor clone was con-sidered fixed. WhenF-ratios were significant, meanseparation was calculated according to SNK method(p < 0.05).

Table 2Analysis of variance and the expected mean square

Source d.f. Expected mean squares

Blocks r − 1 –Treatments t − 1 –Clones (c) c − 1 σ2

e + rσ2cy + ryσ2

cYears (y) y − 1 σ2

e + rσ2cy + rcσ2

yc × y (c − 1)(y − 1) σ2

e + rσ2cy

Error (t − 1)(r − 1) σ2e

r, c andy symbolise numbers of replicates, clones and years, respec-tively; σ2

e: environmental variance;σ2y : variance of year;σ2

c : variance

of clones;σ2cy: variance due toc × y interaction effect.

2.2.2. Phenotypic and genotypic variancesIn order to estimate variance components, the

ANOVA was carried out according to a factorial ran-dom design with years and clones as random factors(Table 2).

The phenotypic variance (σ2p) of a character com-

prises the genotypic variance (σ2g) and the environmen-

tal variance (σ2e). This relationship could be expressed

symbolically as follows:

σ2p = σ2

g + σ2e

whereσ2e andσ2

g were estimated using EMS as follows:

EMSclones− EMSclones× year = ryσ2g

(σ2e + rσ2

cy + ryσ2c ) − (σ2

e + rσ2cy) = ryσ2

c = ryσ2g,

wherer are replicates number andy = 2; thenσ2g could

be computed.Sinceσ2

e = EMS for error, phenotypic variance wascomputed as follows:

σ2p = σ2

g + EMS.

In this study, the broad sense heritability (h2) wasestimated calculating the ratio of genotypic variance(σ2

g) and phenotypic variance (σ2p). The genotypic coef-

ficient of variation (gcv) and phenotypic coefficient ofvariation (pcv) were computed for each trait, in orderte

g

o make comparisons among different traits (Johnsont al., 1955), as follows:

cv =√

σ2g

x100 and pcv=

√σ2

p

x100.

216 S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222

3. Results

Data recorded on daily minimum and maximumtemperatures and rainfall were typical of the Mediter-ranean environment (Fig. 2). In both years, along thegrowing season the minimum temperature increasedlinearly from 8◦C in May to almost 18–20◦C (in Julyand August); it decreased during the winter seasonin the first year till around 6◦C, while in the secondyear the autumn–winter values reached around 4◦Cbetween November and February.

The maximum temperature increased, in 1997, from15◦C in May till 35◦C in June and July, while in 1998,temperatures attained at values higher than 40◦C, neverlower than 30◦C. In the autumn–winter period, in bothyears, minimum temperatures never decreased below0◦C with an average of 5–6◦C.

In the period before rhizome transplanting (fromAugust 1996 until April 1997) 395.3 mm of rain-fall were recorded; during winter periods rainfallsof 385.8 and 128.8 mm were recorded, respectively,in 1997/1998 and 1998/1999. Dry periods wereobserved between May and middle of August in bothyears.

In the first year, the sprouting was affected by thetransplanting date and occurred between April the 20thand May the 18th, after 26 days from transplant. In1998, plant sprouting occurred in all clones betweenMarch the 29th and April the 2nd, which corresponds,

generally, to the period when the plant is sprouting inSouthern Europe environments (Table 3).

In 1997, flowering was observed from Septemberthe 17th till the 21st with the exception of two clones:2 and 7, which did not presented panicle. The year after,with the exception of clones 13, 27 and 39, the majorpart of the clones flowered between September the 26thand October the 10th. Two clones did not flower as well(clones 18 and 19).

The morphological characters measured in the yearsare reported inTables 4 and 5. With reference to thefirst year, all the studied characters, with the exceptionof the medium diameter of the stems, showed signif-icant differences. In the second year, when the fullestablishment of the rhizomes was definitively com-pleted, the characters showing significant differenceswere reduced to six (stem height, plant nodes, stemdensity, base thickness, stem weight and yield).

Average plant height values were of 318.2 and339.4 cm, respectively, in the first and in the secondyear. The clones showed a high variability within thischaracter in both years: in the first year clones 19 andclone 12 showed the highest plant height (333.7 and330.0 cm), respectively. The lowest plant height wasobserved in a wide group with 14 clones with a heightranging from 267.5 to 232.5 cm. In the second year,the range was similarly wide with a maximum valueof 414.3 cm in clone no. 3 and a minimum value of313 cm in clones no. 33, 35, 8, 28.

iment a

Fig. 2. Meteorological conditions during the field exper t the Faculty of Agriculture station in Catania till 1996 to 1999.

S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222 217

Table 3Dates of rhizome sprouting and flowering in both years

Clones 1997 1998

Sprouting Flowering Sprouting Flowering

1 23/4 18/9 30/3 26/92 15/5 No 31/3 26/93 15/5 17/9 31/3 26/94 12/5 18/9 31/3 26/95 23/4 20/9 31/3 26/96 30/4 20/9 31/3 26/97 20/4 No 31/3 10/108 25/4 18/9 31/3 10/109 20/4 20/9 31/3 10/10

10 25/4 20/9 31/3 26/911 25/4 18/9 31/3 26/912 15/5 20/9 31/3 13/913 23/4 18/9 30/3 29/914 25/4 20/9 2/4 10/1015 20/4 19/9 31/3 10/1016 12/5 20/9 30/3 10/1017 1/5 21/9 30/3 10/1018 8/5 18/9 29/3 No19 7/5 18/9 30/3 No20 7/5 20/9 30/3 10/1021 13/5 20/9 31/3 10/1022 10/5 20/9 29/3 10/1023 10/5 21/9 29/3 29/924 2/5 20/9 30/3 29/925 12/5 20/9 31/3 29/926 10/5 18/9 30/3 29/927 8/5 19/9 1/4 21/1028 10/5 19/9 1/4 10/1029 10/5 18/9 2/4 29/930 18/5 20/9 31/3 10/1031 15/5 20/9 30/3 29/932 11/5 20/9 30/3 10/1033 12/5 20/9 30/3 29/934 15/5 19/9 31/3 10/1035 15/5 19/9 31/3 29/936 12/5 18/9 31/3 29/937 15/5 19/9 31/3 29/938 12/5 19/9 31/3 29/939 15/5 20/9 29/3 10/9

The average number of nodes per stem was, in 1997,of 33.0 ranging from 37.1 (clones 1 and 20) and 28.6(clone 29) and in 1998 equal to 33.9 ranging from 38.9(clone 6) to 28.7 (clone 28).

The overall number of stems per square meter(stem density) recorded at the end of the growingseason, doubled, in the average, from the first year(10.4 stems m−2) to the second one (21.4 stems m−2).The variability measured in the second year was

Fig. 3. Relation between aboveground biomass of clones in the firstand second year. For the correspondence between number and nameseeTable 1.

much wide for this character with a difference of 13stems m−2 between the higher value (29 stems m−2 ofclones 35, 19 and 20) and the lowest one (16.2 stemsm−2 of clone 8).

With reference to stem weight, important characteras the previous ones for yield composition, the aver-age value was 141.0 and 127.6 g in the first and in thesecond year, respectively. The variability among clonesranged from 89.1 to 188.7 g in the first year and from103.1 to 164.3 g in the second one.

Above ground biomass yield of the 39 clones stud-ied was, in the average, 10.6 t ha−1 of dry matter in thefirst year and 22.1 t ha−1 in the second one. The clone 4(Piazza Armerina) and the clone 20 (Capo d’Orlando)maintained their high productive aptitude in both years;they yielded, respectively, 13.1 and 14.1 t ha−1 in thefirst year and 34.2 and 26.9 t ha−1 in the second one.The lowest yield, equal to 7.3 t ha−1 (clone 8) was notsignificantly different from those of more of 50% ofclones in the collection. In the second year, a sim-ilar behaviour with a minimum value of 14.9 t ha−1

(clone 32). The uniformity of the yield in the 2 years isunderlined by the positive and significant value of thecoefficient of correlation of the biomass yield in the 2years (r = 0.50** ) (Fig. 3).

218S.L

.Cosentino

etal./IndustrialCrops

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roducts23

(2006)212–222

Table 4Morphological characters and yield of the 39 clones at harvest in the first year (1997/1998)

Clones Stem height(cm)

Plant nodes(n)

Stem density(n m−2)

Base diameter(mm)

Mediumdiameter(mm)

Top diameter(mm)

Base thickness(mm)

Mediumthickness(mm)

Apicalthickness(mm)

Stems (g) Yield(t ha−1)

1 270.0 bh 37.2 a 10.7 ai 16.0 ac 12.3 a 7.25 b 0.82 b 0.73 b 0.55 bd 137.8 ae 10.6 ah2 260.0 ci 33.4 ab 8.7 gi 16.5 ac 14.3 a 9.00 ab 0.97 ab 0.95 ab 0.70 ad 157.0 ac 9.7 bh3 280.0 bf 32.6 ab 8.4 hi 17.5 ac 14.5 a 9.25 ab 0.97 ab 1.10 ab 0.70 ad 136.1 ae 8.7 fh4 293.0 bd 30.9 ab 12.7 ab 20.5 a 16.0 a 11.0 ab 1.12 ab 1.20 a 0.80 ad 184.8 ab 13.1 ac5 250.0 ei 33.1 ab 10.2 bi 19.5 ac 15.5 a 10.5 ab 1.10 ab 1.08 ab 0.75 ad 160.1 ac 10.2 bh6 264.0 bi 30.8 ab 11.8 af 18.5 ac 17.0 a 9.00 ab 1.05 ab 1.10 ab 0.66 ad 154.9 ad 11.4 ag7 242.0 fi 33.6 ab 11.7 ae 14.3 bc 12.8 a 8.00 b 0.90 ab 0.98 ab 0.60 ad 93.2 de 9.2 dh8 249.5 ei 35.1 ab 9.1 fi 18.0 ac 12.8 a 7.00 b 0.90 ab 0.85 ab 0.48 cd 113.3 ce 7.3 h9 238.5 gi 32.0 ab 11.5 ag 18.5 ac 14.3 a 9.50 ab 1.00 ab 1.08 ab 0.63 ad 119.4 ce 9.2 dh

10 294.5 bd 32.7 ab 13.3 a 17.5 ac 15.8 a 8.00 b 1.00 ab 1.05 ab 0.60 ad 151.3 ad 10.2 bh11 284.0 be 30.6 ab 12.8 ab 18.3 ac 15.3 a 8.25 ab 1.07 ab 1.03 ab 0.55 bd 133.0 ae 13.3 ab12 330.0 a 29.6 ab 11.7 ag 18.0 ac 16.3 a 10.0 ab 1.00 ab 1.03 ab 0.70 ad 188.7 a 10.5 ah13 275.0 bg 31.2 ab 10.5 bi 15.3 ac 12.3 a 8.00 b 0.87 ab 0.92 ab 0.75 ad 139.4 ae 12.6 af14 280.0 bf 30.5 ab 10.9 ai 20.8 a 17.3 a 9.50 ab 1.20 a 1.22 a 0.55 bd 172.1 ac 11.8 ag15 285.0 be 31.1 ab 9.8 ci 18.3 ac 14.5a 8.75 ab 1.02 ab 0.98 ab 0.67 ad 147.8 ae 12.2 ag16 304.0 b 32.9 ab 10.3 bi 17.3 ac 16.0 a 10.8 ab 1.07 ab 0.98 ab 0.88 ab 146.0 ae 10.2 bh17 264.3 bi 33.8 ab 9.1 fi 16.8 ac 14.0 a 9.75 ab 0.97 ab 1.05 ab 0.70 ad 133.5 ae 10.2 bh18 275.5 bg 35.8 ab 9.5 di 18.0 ac 13.5 a 9.00 ab 0.85 ab 1.00 ab 0.58 ad 120.2 ce 11.0 ah19 333.67 a 35.1 ab 12.7 ac 19.0 ac 14.8 a 7.25 b 0.92 ab 0.98 ab 0.45 d 121.1 ce 11.6 ag20 284.0 be 37.0 a 12.7 ac 16.3 ac 13.5 a 8.00 b 0.90 ab 0.90 ab 0.60 ad 142.3 ae 14.0 a21 275.0 bg 35.7 ab 9.5 di 16.0 ac 13.0 a 9.75 ab 0.87 ab 1.05 ab 0.85 ac 163.4 ac 12.0 ag22 294.5 bd 34.0 ab 8.1 i 18.8 ac 17.0 a 10.3 ab 1.17 ab 1.15 ab 0.60 ad 172.0 ac 10.2 bh23 284.0 be 36.3 ab 9.4 di 19.3 ac 15.3 a 9.25 ab 1.10 ab 0.92 ab 0.58 ad 136.4 ae 10.2 bh24 293.0 bd 33.6 ab 8.9 fi 19.0 ac 15.5 a 10.0 ab 1.07 ab 1.13 ab 0.58 ad 160.7 ac 9.1 eh25 280.0 bf 34.3 ab 10.6 ai 20.3 ab 15.3 a 10.5 ab 1.07 ab 1.13 ab 0.67 ad 158.6 ac 11.2 ag26 300.0 bc 35.1 ab 10.3 bi 18.3 ac 15.5 a 9.00 ab 0.95 ab 1.15 ab 0.67 ad 142.4 ae 12.8 ae27 286.7 be 34.2 ab 9.4 di 20.3 ab 15.0 a 12.3 a 1.10 ab 1.15 ab 0.95 a 171.6 ac 13.0 ad28 267.0 bi 32.5 ab 12.3 ad 17.3 ac 14.5 a 9.00 ab 0.92 ab 0.92 ab 0.63 ad 122.7 be 10.8 ah29 277.5 bf 28.6 b 10.2 bi 16.8 ac 14.0 a 10.0 ab 1.02 ab 1.08 ab 0.88 ab 141.2 ae 9.3 ch30 276.5 bg 31.0 ab 9.4 di 18.0 ac 16.5 a 10.5 ab 1.05 ab 1.13 ab 0.80 ad 145.3 ae 11.1 ah31 257.5 di 30.4 ab 10.3 bi 14.0 c 11.5 a 8.00 b 0.97 ab 0.90 ab 0.58 ad 120.1 ce 9.0 eh32 233.6 hi 29.0 ab 10.1 bi 19.8 ac 12.3 a 10.0 ab 0.97 ab 1.20 a 0.73 ad 157.5 ac 9.5 bh33 275.0 bg 31.2 ab 9.4 di 18.0 ac 12.3 a 8.50 ab 1.05 ab 0.92 ab 0.70 ad 126.7 ae 8.8 fh34 232.5 i 31.6 ab 11.2 ah 20.0 ac 11.5 a 9.25 ab 1.07 ab 0.98 ab 0.70 ad 116.6 ce 9.6 bh35 267.5 bi 31.2 ab 12.2 ae 15.8 ac 12.3a 8.50 ab 0.97 ab 0.90 ab 0.53 bd 89.1 e 10.6 ah36 283.8 be 34.4 ab 9.3 ei 19.3 ac 14.5 a 8.50 ab 1.00 ab 1.05 ab 0.58 ad 112.5 ce 8.8 fh37 285.7 be 35.9 ab 8.2 hi 19.3 ac 17.5 a 9.75 ab 1.05 ab 1.05 ab 0.75 ad 146.7 ae 8.3 gh38 250.4 ei 34.0 ab 9.0 fi 18.0 ac 13.3 a 10.8 ab 0.92 ab 1.00 ab 0.90 ab 129.8 ae 8.7 fh39 246.5 ei 35.5 ab 10.1 bi 17.0 ac 14.3 a 9.00 ab 0.90 ab 0.98 ab 0.73 ad 132.2 ae 12.1 ag

Average 275.0 33.01 10.4 17.9 14.4 9.24 1.00 1.02 0.67 141.0 10.6

Values with different letters are significantly different atp ≤ 0.05, according to SNK.

S.L.C

osentinoetal./IndustrialC

ropsand

Products

23(2006)

212–222219

Table 5Morphological characters and yield of the 39 clones at harvest in the second year (1998/1999)

Clones Stem height(cm)

Plant nodes(n)

Stem density(n m−2)

Base diameter(mm)

Mediumdiameter(mm)

Top diameter(mm)

Base thickness(mm)

Mediumthickness(mm)

Apicalthickness(mm)

Stems(g)

Yield(t ha−1)

1 320.7 gm 32.4 ac 19.9 fj 20.3 a 17.0 a 7.5 a 1.18 ab 1.28 a 0.43 a 129.0 ad 20.6 be2 367.9 bf 34.1 ac 17.8 ik 17.3 a 15.5 a 7.3 a 1.10 ab 1.13 a 0.45 a 134.7 ad 22.0 be3 414.3 a 33.8 ac 17.9 ik 18.3 a 15.3 a 7.5 a 1.13 ab 1.08 a 0.45 a 112.0 d 21.5 be4 345.0 ck 36.7 ac 27.3 ab 19.0 a 15.3 a 8.0 a 1.10 ab 1.15 a 0.53 a 157.1 ac 34.2 a5 392.0 ab 34.0 ac 21.2 di 18.0 a 14.8 a 7.0 a 1.07 ab 1.05 a 0.43 a 122.5 bd 25.5 bc6 364.0 bg 38.9 a 21.7 di 18.8 a 15.8 a 7.0 a 1.07 ab 1.15 a 0.43 a 139.3 ad 23.6 bd7 338.0 dl 33.7 ac 21.1 di 17.3 a 14.5 a 6.8 a 1.05 ab 1.08 a 0.43 a 105.2 d 18.0 ce8 313.5 im 35.9 ac 16.2 k 19.0 a 14.5 a 7.0 a 1.15 ab 1.15 a 0.38 a 138.5 ad 16.7 de9 356.0 bj 29.7 bc 20.0 ej 18.0 a 14.0 a 6.5 a 1.05 ab 1.05 a 0.38 a 109.1 d 20.5 be

10 346.7 cj 32.7 ac 23.1 cg 18.5 a 15.3 a 5.8 a 1.10 ab 1.05 a 0.33 a 116.2 cd 22.0 be11 381.7 ad 36.8 ac 19.7 fj 18.5 a 16.5 a 6.3 a 0.95 b 1.13 a 0.40 a 132.9 ad 23.2 be12 380.5 ad 34.6 ac 20.7 di 17.5 a 15.3 a 6.8 a 1.02 ab 1.03 a 0.40 a 164.3 a 26.4 bc13 364.9 bg 35.4 ac 21.1 di 18.3 a 14.8 a 6.0 a 1.05 ab 1.08 a 0.35 a 121.5 bd 23.6 bd14 361.5 bh 35.6 ac 19.5 gk 19.3 a 15.5 a 6.8 a 1.15 ab 1.13 a 0.45 a 159.3 ab 24.2 bd15 342.7 ck 32.6 ac 21.1 di 20.0 a 16.3 a 6.0 a 1.13 ab 1.25 a 0.38 a 134.7 ad 22.4 be16 342.3 ck 31.5 ac 19.1 hk 18.0 a 16.0 a 6.8 a 1.13 ab 1.15 a 0.43 a 120.3 bd 22.1 be17 325.0 fm 35.8 ac 16.1 k 19.8 a 15.0 a 7.0 a 1.13 ab 1.08 a 0.45 a 133.3 ad 16.6 de18 375.0 be 37.8 ab 17.8 ik 20.0 a 16.5 a 6.3 a 1.17 ab 1.20 a 0.38 a 145.8 ad 20.7 be19 387.4 ac 32.1 ac 29.5 a 15.8 a 12.5 a 5.5 a 0.97 ab 0.93 a 0.35 a 103.1 d 24.1 bd20 365.2 bg 34.7 ac 29.4 a 17.5 a 14.8 a 6.0 a 1.07 ab 1.03 a 0.35 a 111.7 d 26.9 b21 365.5 bg 36.1 ac 23.5 cf 21.5 a 15.0 a 6.5 a 1.18 ab 1.08 a 0.40 a 116.4 cd 22.0 be22 361.8 bh 30.0 bc 18.9 hk 19.0 a 16.0 a 5.4 a 1.13 ab 1.13 a 0.40 a 141.7 ad 22.7 be23 364.3 bg 36.5 ac 21.1 di 20.0 a 16.8 a 7.0 a 1.07 ab 1.18 a 0.40 a 158.7 ab 20.3 be24 334.0 el 36.2 ac 16.6 jk 18.5 a 15.5 a 6.5 a 1.07 a 1.13 a 0.40 a 134.6 ad 22.1 be25 334.0 el 34.7 ac 19.8 fj 17.5 a 15.3 a 6.3 a 1.13 ab 1.13 a 0.38 a 130.9 ad 19.4 be26 357.3 bi 33.8 ac 21.4 di 18.5 a 15.0 a 6.5 a 1.10 ab 1.18 a 0.38 a 133. 9 ad 23.8 bd27 346.5 cj 32.6 ac 18.6 hk 19.8 a 16.3 a 6.3 a 1.20 ab 1.20 a 0.40 a 133.2 ad 22.5 be28 310.6 jm 28.7 c 24.9 bc 17.5 a 14.5 a 6.5 a 1.45 a 1.18 a 0.53 a 126.4 ad 23.3 be29 329.2 em 30.6 bc 22.1 ch 18.3 a 15.3 a 6.5 a 1.07 ab 1.05 a 0.33 a 107.8 d 21.0 be30 344.3 ck 33.4 ac 21.6 di 19.8 a 16.8 a 6.8 a 1.18 ab 1.20 a 0.43 a 125.3 ad 20.4 be31 318.3 hm 30.7 bc 19.5 gk 20.3 a 16.5 a 7.0 a 1.20 ab 1.18 a 0.43 a 104.8 d 21.9 be32 297.4 ln 35.8 ac 23.9 cd 20.0 a 15.8 a 5.8 a 1.15 ab 1.18 a 0.40 a 119.0 bd 14.9 e33 314.5 im 32.7 ac 21.3 di 17.0 a 14.8 a 6.3 a 1.00 ab 1.03 a 0.38 a 114.5 d 22.6 be34 338.3 dl 35.3 ac 27.3 ab 20.0 a 15.3 a 6.0 a 1.18 ab 1.00 a 0.43 a 116.3 cd 19.3 be35 314.3 im 34.5 ac 28.7 a 19.0 a 15.3 a 6.5 a 1.07 ab 1.13 a 0.40 a 123.9 ad 22.8 be36 301.0 kn 33.4 ac 22.9 cg 19.5 a 15.3 a 7.00 a 1.13 ab 1.03 a 0.43 a 125.7 ad 21.4 be37 345.8 cj 36.4 ac 19.0 hk 20.5 a 16.0 a 6. 0 a 1.25 ab 1.25 a 0.38 a 129.5 ad 26.0 bc38 290.0 mn 30.5 bc 23.7 cd 19.3 a 14.3 a 6.3 a 1.20 ab 1.18 a 0.43 a 110.9 d 19.7 be39 269.3 n 32.5 ac 20.4 di 19.8 a 16.0 a 7.0 a 1.20 ab 1.23 a 0.40 a 131.1 ad 21.7 be

Average 339.4 33.9 21.4 18.8 15.4 6.55 1.12 1.12 0.41 127.6 22.1

Values with different letters are significantly different atp ≤ 0.05, according to SNK.

220 S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222

Table 6Analysis of variance—mean squares for analysis of 11 characters combined inArundo donax (significance of years and clones was calculatedwith respect to “year× clone” interaction)

Character Mean squares

Year Clones Year× clones

Yield (t ha−1) 5195.32** 18.23** 7.75Stem weight (g) 7012.20** 1119.33** 383.45Stem density (n m−2) 4721.93** 19.85** 7.63Base thickness (mm) 1.19** 0.02 0.04*

Medium thickness (mm) 0.73** 0.03 0.04Apical thickness (mm) 5.63** 0.03 0.03***

Stem nodes (n) 32.68** 11.32 9.98Base diameter (mm) 63.72** 8.49 7.88Medium diameter (mm) 70.21** 7.92 6.02Apical diameter (mm) 552.00** 3.44 2.90**

Stem height (cm) 466355.80** 5094.05** 2079.55

* Significant at 5% probability level.** Significant at 1% probability level.

*** Significant at 0.1% probability level.

3.1. Phenotypic and genotypic variability

The mean squares for year, clones and year× clonesinteraction with respect to the eleven characters stud-ied are present inTable 6. Four characters: biomassyield, stem weight, stem density and height of stemsshowed a significant variance among clones. Interac-tions “year× clone” are significant for base thickness,apical thickness and apical diameter. The others did notshow any statistical significance.

3.2. Phenotypic and genotypic coefficients ofvariation (pcv and gcv) and heritability (h2)

The genetic analysis made for the four characterswhich showed stable differences among clones, is pre-sented inTable 7. The gcv, pcv andh2 values are

presented for the characters investigated. In both years,the range of phenotypic values in comparison to thegeneral mean showed wide variability for most of thecharacters studied (Table 7).

The selected characters showed moderate values ofheritability: yield 23%, stem weight 25%, stem densityand stem height 48 and 46%, respectively.

3.3. Phenotypic correlations

The phenotypic correlations are reported inTable 8for the first and the second year. Yield was posi-tively and significantly correlated in both years tostem weight (r = 0.35* and 0.33* in the first and sec-ond, respectively), number of stem per square meter(r = 0.46** and 0.59*** ) and plant height (r = 0.34* and0.35* ).

Table 7Phenotypic and genotypic components of variability, coefficients of variation and broad sense heritability in characters ofArundo donax computedin both years

Character Value Variance Coefficient Heritability

Mean Range Phenotypic Genotypic pcva (%) gcvb (%) h2 (%)

Yield (t ha−1) 16.30 7.3–34.2 11.42 2.62 20.67 9.90 23Stem weight (g) 134.3 89.1–188.7 747.87 183.97 20.37 10.10 25Stem density (n m−2) 15.9 8.1–29.5 6.43 3.06 15.92 10.98 48Stem height (cm) 309.6 232.5–414.3 658.79 301.45 8.29 5.61 46

a Phenotypic coefficient of variation.b Genotypic coefficient of variation.

S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222 221

Table 8Correlation matrix (r) among four characters computed in both years

X1 X2 X3

I yearYield (t ha−1) (X1) –Stem weight (X2) 0.35*

Stem density (X3) 0.46** −0.11Stem height (X4) 0.34* 0.41** 0.14

II yearYield (t ha−1)Stem weight (X1) 0.33*

Stem density (X2) 0.59*** −0.29Stem height (X3) 0.35* 0.16 −0.07

* Significant at 5% probability level.** Significant at 1% probability level.

*** Significant at 0.1% probability level.

The multiple regression analysis performed takinginto consideration yield as dependent variable and stemweight, stem density and stem height as independentvariables gave more indications on the effect of yieldcomponents on yield. In both years, the results showthat stem density has the highestF values (F = 14.6***

and 63.12*** ) and then seems to have the most impor-tant influence on yield. Stem weight, instead, shows alow F value in the first years (F = 7.1* ), whereas it ishigher in the second one (F = 13.3*** ). Height does notaffect yield in the first year (F = 0.82 ns), but in the sec-ond year this character is significantly related to yield(F = 13.0) (Table 9).

Table 9Partial coefficients of the multiple regression for stem weight, stemsdensity and stem height vs. yield in both years

MS coefficient F

I yearIntercept 7.517***

Stem weight 0.0245 7.112*

Stem density 0.5364 14.615***

Stem height 0.0093 0.8231 ns

II yearIntercept 29.779***

Stem weight 0.1048 13.325***

Stem density 0.7215 63.120***

Stem height 0.0353 12.892**

* Significant at 5% probability level.** Significant at 1% probability level.

*

4. Discussion and conclusions

The results of this study carried out with the objec-tive to individuate a variability in the germplasm forbreeding programme inA. donax L. in areas of theSouth Italy, and in particularly in Sicily, could givesome answer to the objective of research which aimedat studying the possibility to undertake breeding workto obtain genotypes ofA. donax L. with a high yieldpotentiality.

The yields obtained in this study attained at verysatisfying values, especially for clone no. 4 (PiazzaArmerina-Fegotto) and clone no. 20 (Capo d’Orlando),which maintained their productive aptitude in bothyears.

These results are comparable to what obtainedin experimental trial carried out in Spain (17.0 and22.5 t ha−1, respectively, in the first and second year)and in South of Greece (13.7 and 19.0 t ha−1, respec-tively in the first and second year) (Christou, 2001).Results of trials carried out by C.E.T.A. (Vecchiet andJodice, 1996) reported yields in the second year rang-ing from 15.7 to 36.8 t ha−1. In our experiment, in thesecond year as well, a similar range of values (from15.0 to 34.2 t ha−1) was observed.

The analysis of variance has pointed out an absenceof significant variability for many characters: base,medium and apical thickness, stem nodes, base,medium and apical diameter.

Moreover, respect to the characters with signifi-c sb ctiono duet very nsid-e nifi-c fulo ofs esec 16%a ilityf saidc4 vet

berp oveg are

** Significant at 0.1% probability level.

ant interaction effects of clone× year, the differenceetween clones varied over years so that selef clones for these characters would be difficult

o the lack of stability of the average effects oears. Therefore, these characters were not cored for genetic improvement. The statistical sigance for main effect ‘clone’ was found to be usenly in four characters (yield, stem weight, densitytems and height of stems). The first three of thharacters had a considerable pcv higher thannd a gcv more than 9% indicating some possib

or genetic improvement. Moreover, the aboveharacters had moderate heritability (h2) (from 23 to8%) showing the possibility of selecting to impro

hem.The positive correlation between the stem num

er m2, the stem weight and the stem height with abround dry biomass yield indicates that those traits

222 S.L. Cosentino et al. / Industrial Crops and Products 23 (2006) 212–222

the most important yield components. Indeed in theabove-mentioned characters, which have an agronomicimportance for biomass production, a wide variability,within the studied clones, was observed.

The stem density resulted highly related to yield, fol-lowed by the weight. It is worth to recall that in researchcarried out using the random amplified polymorphicDNA (RAPD) method, was observed a reduced vari-ability caused by the common origin of the clones. Thisobservation indicates the low genetic polymorphismamong the studied populations and a rather uniformgenetic pool (Lewandowski et al., 2003). However,according to the heritability observed in this study, itseems that may exists a genetic base for these charac-ters among these clones, which should be better studiedand could be fruitfully exploited.

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