Pinus Roxburghii Sarg Research Articles

Floristics diversity composition along an altitudinal gradient in Himalayan chir pine forest of the western Himalayas

In the present study, we examined the species composition of Himalayan Chir Pine forest (Pinus roxburghii Sargent) at different altitudes between 1000-1600m asl, considering three altitudinal gradients viz., A1(Lower altitude: 1000-1200m); A2(Middle altitude: 1200-1400m); A3(Higher altitude: 1400-1600m) in the Western Himalayas. In this study, a total of 50 species were recorded, of which 9 were trees, 15 shrubs, 26 herbs. Herb plant richness (in number) showed a diminishing trend with altitude whereas shrub species remain same for low and mid altitudes with slight drop in high altitudes and the tree species richness remain constant for all the altitudes. Many types of environmental changes (altitudes, slopes, aspect, temperature, precipitation etc.) influence the processes that can both augment or erode diversity. The study suggests that species distribution, presence of the particular species (species composition) and number of species present (species richness) are largely regulated by altitudes and other climatic + topographic factors.

Pinus Roxburghii Sarg. Ameliorates Alzheimer?s disease- Type Neurodegeneration and Cognitive Deficits Caused by Intracerebroventricular-Streptozotocin in Rats: An In Vitro and In Vivo Study

Pinus roxburghii Sarg. is traditionally used as a nerve tonic, hemostatic and anticonvulsant. The present study was aimed to investigate the neuroprotective activity of Pinus roxburghiiSarg. stem bark for memory and cognition using in vitro and in vivo methods. Plant drug was extracted with methanol using Soxhlet extraction. In vitro cell viability activity of Pinus roxburghiiSarg. was assessed using PC-12 cell lines. The in vivo neuroprotective activity of Pinus roxburghii Sarg. was tested on Wistar albino rats (either sex) and intracerebroventricular streptozotocin (3 mg/kg, bilateral) was given to cause memory deficits. High-performance thin layer chromatography fingerprinting was performed for the presence of quercetin and gallic acid in the plant extract. The phytochemical testing of extract revealed the presence of flavonoids, alkaloids, steroids, and tannins. Pinus roxburghii Sarg. showed significant cell viability at 10, 50 and 100 μg/ml in an in vitro test on PC-12 cell lines. In the in vivo activity, intracerebroventricular streptozotocin caused significant deterioration in memory and cognition, tissue oxidative stress and acetylcholinesterase activity. Pinus roxburghiiSarg. 100, 200 and 300 mg/kg, per os and donepezil (3 mg/kg, per os) significantly (p<0.05) reversed the behavioral changes in rats when tested on morris water maze and elevated plus maze. Increased levels of lipid peroxidation, acetylcholinesterase activity and decreased the level of glutathione were significantly (p<0.05) antagonized by Pinus roxburghii Sarg., similar to donepezil in rat brain. Quercetin and gallic acid were identified quantitatively in the plant extract in High-performance thin layer chromatography study. The methanolic extract of Pinus roxburghii Sarg. ameliorated intracerebroventricular streptozotocin induced cognitive deficits and memory impairment possibly due to the presence of quercetin and gallic acid.

Chemical constituents of the extracts of the knotwood of Pinus roxburghii Sarg. and their antioxidant activity

Chemical composition of the extract from the Pinus roxburghii Sarg. knotwood was studied for the first time. The extracts were obtained by ultrasound assisted sequential extraction of knotwood with hexane, acetone—water (95 : 5, v/v), and water. According to GC-MS analysis, hexane extract was dominated by monoterpenoids amounting to 53.57% and sesquiterpenoids (41.05%). Components of the acetone—water (95 : 5, v/v) extract were isolated, purified by HPLC, and identified by NMR spectroscopy and mass spectrometry as 2,3-bis(4-hydroxy-3-methoxybenzyl)butan-1,4-diol, 3-hydroxy-3,4-bis(4-hydroxy-3-methoxybenzyl)dihydrofuran-2(3H)-one and 3,4-bis(4-hydroxy-3-methoxybenzyl)dihydrofuran-2(3H)-one. These compounds demonstrated notable antioxidant capacity determined using the amperometric method.

Estimation of Biomass and Soil Carbon Stock in Pinus roxburghii and Quercus leucotrichophora Forests of District Shimla, Himachal Pradesh

A study was conducted to estimate the biomass and soil carbon stock in various sites of Pinus roxburghii Sarg. (Chir pine) and Quercus leucotrichophora A. Camus (Ban oak) forests of district Shimla, Himachal Pradesh during the year 2015-2016. In case of chir pine forests, the amount of carbon stock in Guma site for above ground, below ground, under storey and litter was 80.05, 16.01, 3.24 and 1.55 tC/ha, respectively. Whereas, values of carbon stock at Dhami site were 192.92, 38.58, 1.71 and 1.57 tC/ha, respectively. The biomass as well as the carbon stock was higher at Dhami site than Guma site. The soil organic carbon stock (tC/ha) at 10-15 cm, 15-30 cm and 30-45 cm was 22.45, 19.99 and 12.36, respectively at Guma site. Whereas, these values at different depths were 23.23, 20.17 and 15.59 tC/ha, respectively for Dhami site. The value of total soil carbon stock was more (58.98 tC/ha) at Dhami site than Guma site (54.80 tC/ha) upto 45cm depth. In case of Ban oak forests, the amount of carbon stock in Taradevi site for above ground, below ground, under storey and litter was 151.27, 37.82, 3.04 and 1.52 tC/ha, respectively. Whereas, values of carbon stock at Koti site were 74.25, 18.56, 1.24 and 1.26 tC/ha, respectively. Total biomass as well as the carbon stock was higher at Taradevi site than Koti site. The soil organic carbon stock (tC/ha) at 10-15 cm, 15-30 cm and 30-45 cm was 42.07, 29.70 and 14.78, respectively at Tardevi site. Whereas, these values for three depths were 36.60 , 27.72 and 18.72 tC/ha, respectively for Koti site. The value of total soil carbon stock was more at Tardevi site (86.55 tC/ha) than Koti site (83.04 tC/ha).

Changes in vegetation cover and soil intrinsic properties influence the soil bacterial community composition and diversity across different climatic regions of India

Soil microbial community in forest ecosystems plays a significant role in carbon and nutrient cycling. Very little is known about the effect of vegetation cover on soil bacterial community composition and diversity across different climatic regions of India. Soil was sampled from the plant cover dominated by seasonal herbs and grasses (PAS), Pinus roxburghii Sarg (PIN), Abies pindrow L. (FIR), Quercus incana Roxb (OKF), Mitragyna parvifolia Roxb (MIK); Acacia nilotica L. and Salvadora spp. (MIA) in three different climatic regions (humid, moist sub-humid and semi-arid) of India. The soil physical, chemical and biological properties such as sand (SAN), silt (SIL), clay (CLY), bulk density (SBD), wilting point (SWP), field capacity (SFC), saturated hydraulic conductivity (SHC), pH (SpH), organic carbon (SOC), nitrogen (SN), C:N ratio, available phosphorous (SAP) and total microbial activity (SMA) were determined. Illumina sequencing of specific 16S rRNA gene was applied to identify bacterial community composition in the soils under different vegetation cover. Results showed that the soil properties varied under different vegetation cover across the different climatic regions. SOC, SN, SMA were highest in the moist sub-humid region sites (PAS, PIN) followed by humid region (OKF, FIR) and semi-arid region (MIK, MIA) sites. However, the Chao 1 (species richness), Shannon and Simpson index (diversity) were highest in OKF, followed by MIK, FIR, MIA, PIN and PAS. The predominant bacterial phyla and genera in the soils under different vegetation cover were Proteobacteria, Actinobacteria, Bacteroidetes, Firmicutes, Planctomycetes, Verrucomicrobia, Thermotogae and Geobacter, Methylocapsa, Sphingomonas, Pseudomonas, Erysipelotrichaceae_incertae_sedis, Sporotomaculum, Amorphus, Helicobacter, Paenibacillus, Bauldia, Skermanella, Methylosinus, Singulisphaera, Marinobacter and Lamprocystis. We also found the exclusive OTUs abundance of some bacterial phyla and genera in the soils, which were not correlated with any one of the studied soil variables. In our analysis, we found only the Firmicutes, Verrucomicrobia and Bacteroidetes are linearly correlated (P < 0.05) with CLY, SWP and SpH. Likewise, bacterial genera Methylocapsa, Methylosinus and Amorphus were linearly correlated (P < 0.05) with SIL, SpH, C:N ratio and SAP. Our results suggested that the type of vegetation cover has a significant impact on changes in soil properties, controlling the soil bacterial community composition and diversity across different climatic regions of India. The soil bacterial community composition and diversity may serve as a potential ecological indicator with respect to land use and land cover change on biogeochemical cycling processes across different climatic regions of India.

Positive relationship between species richness and aboveground biomass in Kumaun Himalayan forest

Understory vegetation is an important part of forest ecosystem which affect the physical and chemical properties of soil, quality and quantity of forest litter and water storage capacity of litter soil layer. Thus, the influence of soil nutrients on the understory species composition of forest ecosystem cannot be ignored. The authors set 15 typical plots with area of 1 × 1 m in chir pine (Pinus roxburghii Sarg.), oak (Quercus leucotrichophora A. Camus)—chir pine mixed and oak–cypress (Cupressus torulosa D. Don) mixed forest. Chemical properties of the soil were analyzed at 0–10 cm, 10–20 cm and 20–30 cm in all the selected forest types. Phytosociological and diversity parameters were also calculated for each forest type. The present study reveals that carbon, nitrogen, pH and phosphorus are the most important environmental drivers which influence understory species composition. Plant species richness significantly correlated with species composition. To maintain the diversity and structure of ecosystems we should consider the co-evolution of both vegetation and soil. Further studies on climate and microorganism are needed to further explore the interactive relationships among vegetation and soil properties.

The Temperate House Restoration Project

There is a long tradition at the Royal Botanic Gardens, Kew (RBG, Kew) of cultivating and displaying exotic plants from all over the globe, and the largest Victorian glasshouse, the Temperate House, traditionally showcases plants from temperate regions. The Temperate House Restoration Project was undertaken at RBG, Kew from 2012 to 2018. Over 1,000 species of plants were removed, propagated and replanted for this project, and this article describes the propagation of some of the most difficult to reproduce plant material. Four plant groups or species are presented: Erica verticillata P.J.Bergius, Quercus insignis M.Martens Galeotti, Pinus roxburghii Sargent and Banksia spp. L. This is in order to illustrate the variety of options available for propagating challenging species with attention to their ecology, biology and growing requirements.&#x0D; Also provided are background information, reasons why these plants are considered difficult to multiply in cultivation, how plant material was sourced and the methods employed which led to successful propagation of the material at RBG, Kew. Propagation of the plants was heavily reliant on the horticultural expertise of those involved, and this expertise ensured that most of the original plant material was rejuvenated and new collections with scientific significance were added to the restored Temperate House.

The Temperate House Restoration Project

There is a long tradition at the Royal Botanic Gardens, Kew (RBG, Kew) of cultivating and displaying exotic plants from all over the globe, and the largest Victorian glasshouse, the Temperate House, traditionally showcases plants from temperate regions. The Temperate House Restoration Project was undertaken at RBG, Kew from 2012 to 2018. Over 1,000 species of plants were removed, propagated and replanted for this project, and this article describes the propagation of some of the most difficult to reproduce plant material. Four plant groups or species are presented: Erica verticillata P.J.Bergius, Quercus insignis M.Martens Galeotti, Pinus roxburghii Sargent and Banksia spp. L. This is in order to illustrate the variety of options available for propagating challenging species with attention to their ecology, biology and growing requirements. Also provided are background information, reasons why these plants are considered difficult to multiply in cultivation, how plant material was sourced and the methods employed which led to successful propagation of the material at RBG, Kew. Propagation of the plants was heavily reliant on the horticultural expertise of those involved, and this expertise ensured that most of the original plant material was rejuvenated and new collections with scientific significance were added to the restored Temperate House.

Exploration of Pinus roxburghii Sargent Dominated Communities in Maidan Valley, Dir (L), Khyber Pakhtunkhwa, Pakistan

Exploration of Pinus roxburghii Sargent Dominated Communities in Maidan Valley, Dir (L), Khyber Pakhtunkhwa, Pakistan

IN VITRO ALPHA-GLUCOSIDASE INHIBITORY ACTIVITY OF EGYPTIAN PLANT EXTRACTS AS AN INDICATION FOR THEIR ANTIDIABETIC ACTIVITY

Objective: Diabetes mellitus is a highly prevalent chronic disease in Egypt leading to high socioeconomic problems, especially in the cities due to the unhealthy life style. Although many drugs are available, they have many side effects. Furthermore, the body arouses resistance after a while for the drug so it should be changed every once in a while. Plants could be a good source for drugs. In Egypt, we have a rich flora which has not been subjected to systematic screening for antidiabetic activity.Methods: The aim of this work was to screen 264 plant extracts for their in vitro α-glucosidase inhibitory activity. Those extracts which gave more than 70% inhibition were screened on different concentrations and their inhibitory concentrations giving 50% activity (IC50) were calculated.Results: Out of all the tested extracts, 63 gave more than or equal 70% inhibition on α-glucosidase at the tested concentration (25 ppm). After the calculation of the IC50 values, 10 extracts were chosen for further study having 5 ppm and less IC50.Conclusion: The most active plant extract is Pinus roxburghii Sarg. branches (IC50 is 2.47 ppm).

Physico-chemical characterization of Oak, Pine and Sal forest soil profiles of Betalghat Region of Kumaun Himalaya

Soil is one of the basic natural resources on which all forms of terrestrial life co-exist. The soils of three forest types i.e., banj-oak ( Quercus leucotrichophora A. Camus) chir-pine ( Pinus roxburghii Sargent) and sal ( Shorea robusta Gaertn. f.) of Betalghat block, Nainital District, (Kumaun Himalaya). Soils were drawn from three depths viz. 0-10 cm, 11-20 cm, 21-30 cm. The present study were studied resulted to soil physico-chemical properties i.e. maximum soil texture (sand 34.30 %, silt 57.21 % and clay 33.23 %), soil moisture ( 12.6 %), bulk density ( 1.43 g.cm -3 ), porosity ( 52.06 %), water holding capacity ( 69.27 %) and the maximum values of chemical analysis i.e. pH ( 6.80 ), total carbon ( 3.82 %), organic matter (6.57 %), total nitrogen ( 0.36 % ), available potassium ( 267 mg/kg ) and available phosphorous ( 62 mg/kg ). Banj-oak forest shows rich physical properties followed by sal and chir-pine forest while chir-pine forest shows rich chemical properties in compared to sal and banj-oak forest.

Spatial distribution of Pinus roxburghii Sarg. (Pinaceae) in Rasuwa District of Nepal with the application of Remote Sensing and Geographic Information System

Spatial distribution of Pinus roxburghii Sarg. (Pinaceae) in Rasuwa District of Nepal with the application of Remote Sensing and Geographic Information System

Moisture-Limited Tree Growth for a Subtropical Himalayan Conifer Forest in Western Nepal

Chir pine (Pinus roxburghii Sarg.) is a common tree species with ecological and economic importance across the subtropical forests of the central Himalayas. However, little is known about its growth response to the recent warming and drying trends observed in this region. Here, we developed a 268-year-long ring-width chronology (1743–2010) from western Nepal to investigate its growth response to climate. Based on nearby available meteorological records, growth was positively correlated with winter (November to February; r = 0.39, p &lt; 0.05) as well as March to April (r = 0.67, p &lt; 0.001) precipitation. Growth also showed a strong positive correlation with the sum of precipitation from November of the previous year to April of the current year (r = 0.65, p &lt; 0.001). In contrast, a negative relationship with the mean temperature in March to April (r = −0.48, p &lt; 0.05) suggests the influence of warming-induced evapotranspiration on tree growth. Spring droughts lasting 4–6 months constrain Chir pine growth. These results are supported by the synchronization between droughts and very narrow or locally missing rings. Warming and drying tendencies during winter and spring will reduce forest growth and resilience and make Chir pine forests more vulnerable and at higher risk of growth decline and dieback.

Variation in structure and composition of two pine forests in Kailash Sacred Landscape, Nepal

Chir pine (Pinus roxburghii Sarg.) and blue pine (Pinus wallichiana A.B. Jacks.) are two common species found in mid-hill forests of Nepal where households largely depend on forest resources for their livelihoods and subsistence. The management of such forests is supported by our understanding of the dynamics in forest structure and species composition and the relationship between different forest community characteristics. This study was designed to determine the variation in species composition and the relationship between various forest community characteristics in two pine forests of Kailash Sacred Landscape, Nepal. Quadrat sampling was applied to collect information on forest species, forest community structure, and disturbance factors.Data was statistically analyzed using IBM SPSS. There were a total of 31 plant species under 28 genera and 20 families in the P. roxburghii forest, and 38 plant species under 37 genera and 19 families in the P. wallichiana forest. Mean DBH, height and canopy diameter of P. roxburghii was 23.98 cm, 12.77 m and 1.97 m, respectively, and that of P. wallichiana was 31.5 cm, 11.48 m and 2.79 m, respectively. The relationship between DBH and both height and crown diameter showed strong relationships in the two forest types. In both forests, DBH and height class distribution showed a hump-shaped (unimodal type) distribution with a greater proportion of medium-sized individuals that indicated disruptive forest regeneration. Fire and tree cut were significant disturbance factors in P. roxburghii forest, while grazing and trampling were significant in P. wallichiana forest. The extent of these disturbance factors as determinants of regeneration and species recruitment is important to assess for effective forest management. Banko JanakariA Journal of Forestry Information for NepalVol. 28, No. 1, 2018, page: 26-36

Germination Response of Ornamental Pine Seeds on a Two-Way Thermogradient Plate

The objective of this study was to evaluate the effect of temperature on germination of Pinus tabuliformis Carriere seed, Pinus gerardiana Wall. ex D. Don and Pinus roxburghii Sarg. Seeds of three species were distributed in Petri dishes with 1% agar solution; for the first step it was placed on a 2-way Grant’s thermogradient plate (a bi-directional incubator) for a period of 28 days and during the second step in an incubator type B.O.D. also for 28 days with non-germinated seeds in the first step. This instrument allows an assessment of a wide regime of constant and alternating temperatures, resulting to 144 different combinations of temperature, scheduled to the temperatures 4°C - 40°C. The lighting regime was 12 hours photoperiod for the two stages of the experiment. The best temperature for the germination of P. tabuliformis was 26.2°C (minimum of 7.9°C and maximum 37.0°C) and P. roxburghii was 15.6°C (minimum 10, 4°C and maximum 37.0°C). The seeds are neutral photoblastic. P. gerardiana showed low germination percentage (below 10%). The temperature that provided the highest germination percentage for P. tabuliformis was 26.2°C and 15.6°C for P. roxburghii.

Temporal variation in tree biomass and carbon stocks of Pinus roxburghii Sargent forests of Rajouri forest division in Jammu &amp; Kashmir State

The present study was conducted on Pinus roxburghii (chir) forests of Rajouri Forest Division in Jammu & Kashmir State to assess the temporal variations of carbon biomass and C storage between 1995-96 and 2015-16. Results of present study unveil that despite an increase of Chir tree density from 132.60 trees/ha during 1995-96 assessment to 197.61 trees/ha during 2015-16, the growing stock as well as aboveground tree biomass of Chir forests, which were 16.28 lac m3 and 9.73 lac tonnes in 1995-96, declined to 12.10 lac m3 and 7.24 lac tonnes, respectively by 2015-16. Similarly, the total C stocks of Chir forests (aboveground + belowground), which stood at 5.90 lac tonnes during 1995-96, declined to 4.39 lac tonnes; a decrease of 25.59% in assessment period. In line with these changes, the C density of these forests also witnessed a decline from 58.95 Mg ha-1 to 43.84 Mg ha-1. Such declining trends definitely indicate forest degradation; poor forest health, cumulatively attributed to excessive removals, skewed distribution of Chir trees, subdued movement of trees from lower to higher diameter classes and increased anthropogenic pressure on forests in the division.

Structural profiling of wax biopolymer from Pinus roxburghii Sarg. needles using spectroscopic methods

Pinus roxburghii Sarg. is the most abundant species in Himalayan region. The needles of the species largely contribute to the forest biomass and remain the major cause of forest fires leading to climate change, biodiversity loss, etc. Intriguingly, the layer of needles contains wax, a biomacromolecule with potential chemical functionalities for value addition. In the present study, a distinctive approach towards complete structural analysis of the isolated wax in its native state has been done using 1H, 13C, HSQC, HMBC, COSY, TOCSY along with GC–MS of the methyl esters of constituent fatty acids. The wax was isolated in a quantitative yield of 1.64% and analyses suggest that it is a polymer of linearly attached fatty acid esters which on hydrolysis yielded three types of ω-hydroxy fatty acids viz. 12-hydroxydodecanoic acid, 14-hydroxytetradecanoic acid and 16-hydroxyhexadecanoic acid in a ratio of 1:1:2 respectively. Complete assignments for a carbonyl group, α-, β- and other methylenes present in wax were achieved; corroborating the presence of polyester. In particular, identification of wax structure was accomplished through NMR; thereby providing a lead towards future structural analysis of waxes in their native form. The study would also be helpful to generate commercially important compounds derived from pine needle wax. This will offer an opportunity for utilisation of pine needle biomass: a root cause of Himalayan forest fires.

Effect of Artificially Induced Drought Stress on Seedlings of Pinus roxburghii Sarg

ABSTRACT Water relation and growth parameters of drought stressed seedlings of Chir pine (Pinus roxburghii Sarg.) were studied over a one year period. The seedlings were subjected to three drought cycles of 9, 18, 27 days and one control level (watered every alternate day) in a polyhouse. Pre-dawn (5.30-6.30 AM) and Mid-day water potential was estimated using a pressure chamber. The bench drying method was followed for developing P-V curves for estimating components of water potential. Osmotic adjustment was estimated as the difference between the osmotic potential at zero and full turgor of seedlings kept at control and those given 27 days drought. Across drought cycles the pre- dawn water potential ranged between -0.59 MPa and -2.12 MPa and the mid-day water potentials were always lower than pre-dawn water potentials. The Osmotic adjustment was – 1.17 MPa at full turgor and -2.0 MPa at zero turgor. The seedlings growth was positively related to soil moisture and water potential. The daily change in water potential (difference between pre-dawn and mid-day water potential) declined with increasing drought and was nominal (less than -0.2 MPa) in seedlings subjected to 18 and 27 days drought cycle. Needle characteristics were severely affected by water stress. In Pinus roxburghii Sarg seedlings osmotic adjustment may be an important adaptive mechanism that assists it to invade and grow on dry inhospitable sites.

Constituents and termiticide potential of some wood extracts against Coptotermes heimi (Wasmann) (Isoptera: Rhinotermitidae)

Wood extractives are one of the main reasons for the resistance of wood to termite attack. A study was carried out to determine the chemical constituents of wood extractives from Pinus roxburghii Sargent , Morus alba L. and Eucalyptus camaldulensis Dehnh against Coptotermes heimi (Wasmann) under laboratory conditions in Forman Christian College University (Lahor, Pakistan) in June 2015. Gas chromatography-mass spectrometry of wood extractives of P. roxburghii detected 1,2 - benzenedicarboxylic acid, mono (2-ethylhexyl) ester and octadecanoic acid (methyl ester). Hexadecanoic acid (methyl ester) and 1-methyl-3-(1-methylethyl)-benzene were present in M. alba in addition to the compounds present in P. roxburghii and in E. camaldulensis 1,2-benzenedicarboxylic acid, mono (2-ethylhexyl) ester and β-phellandrene were also present. Based on the feeding activity, wood extracts were arranged in descending order of preference; P. roxburghii > M. alba >E. camaldulensis . Extracts of P. roxburghii, M. alba and E. camaldulensis proved repellent at higher concentrations, with tunneling activity almost fully inhibited. So these could prove useful in developing a soil barrier to block termite activity and serve as a replacement to synthetic chemicals.

Variability studies for needle and wood traits of different half sib progenies of Pinus roxburghii Sargent.

Genetic variability studies for needle and wood traits were carried out for the different half sib progenies of Chir pine, raised in 1985 at the main campus of University. There existed a significant variation for these traits among the different half sib progenies, viz., needle length (18.1-24.6cm), needle thickness (0.53-0.71mm), number of stomata per mm of a row (7.3-12.0), specific gravity of wood (0.36-0.46), tracheid length (1.51-1.85) and moisture content of wood (47.76-58.81). This variability was found under genetic control, as all these progenies are growing under same environment, and are of same age. Traits having high heritability and genetic gain like, needle thickness, wood specific gravity, tracheid length and others, indicate high genetic control. This variability can be exploited in tree improvement programs through selection and breeding approaches for development of advanced generations. Correlation studies for different traits at genotypic and phenotypic levels provided the basic knowledge of association to chalk out efficient breeding strategy for higher productivity through indirect selection.