The Pinaceae, pine family, are trees or shrubs, including many of the well-known conifer
s of commercial importance such as cedars
s and spruce
s. The family is included in the order Pinales
, formerly known as Coniferales
. Pinaceae are supported as monophyletic by their protein-type sieve cell plastids, pattern of proembryogeny, and lack of bioflavonoids. They are the largest extant conifer family in species diversity, with between 220 and 250 species (depending on taxonomic
opinion) in 11 genera,
and the second-largest (after Cupressaceae
) in geographical range, found in most of the Northern Hemisphere
, with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family often forms the dominant component of boreal, coastal, and montane forests. One species, ''Pinus merkusii
'', grows just south of the equator
in Southeast Asia. Major centres of diversity
are found in the mountains of southwest China
, central Japan
, and California
Members of the family Pinaceae are tree
s (rarely shrub
s) growing from tall, mostly evergreen
(except the deciduous
''Larix'' and ''Pseudolarix''), resinous, monoecious, with subopposite or whorled branches, and spirally arranged, linear (needle-like) leaves.
The embryos of Pinaceae have three to 24 cotyledon
The female cones
are large and usually woody, long, with numerous spirally arranged scales, and two winged seed
s on each scale. The male cones are small, long, and fall soon after pollination; pollen dispersal is by wind. Seed dispersal is mostly by wind, but some species have large seeds with reduced wings, and are dispersed by bird
s. Analysis of Pinaceae cones reveals how selective pressure has shaped the evolution of variable cone size and function throughout the family. Variation in cone size in the family has likely resulted from the variation of seed dispersal mechanisms available in their environments over time. All Pinaceae with seeds weighing less than 90 mg are seemingly adapted for wind dispersal. Pines having seeds larger than 100 mg are more likely to have benefited from adaptations that promote animal dispersal, particularly by birds. Pinaceae that persist in areas where tree squirrels are abundant do not seem to have evolved adaptations for bird dispersal.
Boreal conifers have many adaptions for winter. The narrow conical shape of northern conifers, and their downward-drooping limbs help them shed snow, and many of them seasonally alter their biochemistry to make them more resistant to freezing, called "hardening".
Classification of the subfamilies and genera of Pinaceae has been subject to debate in the past. Pinaceae ecology, morphology, and history have all been used as the basis for methods of analyses of the family. An 1891 publication divided the family into two subfamilies, using the number and position of resin canals in the primary vascular region of the young taproot as the primary consideration. In a 1910 publication, the family was divided into two tribes based on the occurrence and type of long–short shoot dimorphism.
A more recent classification divided the subfamilies and genera based on the consideration of features of ovulate cone anatomy among extant and fossil members of the family. Below is an example of how the morphology has been used to classify Pinaceae.
The 11 genera are grouped into four subfamilies, based on the microscopical anatomy and the morphology of the cones, pollen, wood, seeds, and leaves:
* Subfamily Pinoideae
''): cones are biennial, rarely triennial, with each year's scale-growth distinct, forming an umbo on each scale, the cone scale base is broad, concealing the seeds fully from abaxial
view, the seed is without resin vesicles, the seed wing holds the seed in a pair of claws, leaves have primary stomatal bands adaxial (above the xylem) or equally on both surfaces.
* Subfamily Piceoideae
''): cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds fully from abaxial view, seed is without resin vesicles, blackish, the seed wing holds the seed loosely in a cup, leaves have primary stomatal bands adaxial (above the xylem) or equally on both surfaces.
* Subfamily Laricoideae
'', and ''Cathaya
''): cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds fully from abaxial view, the seed is without resin vesicles, whitish, the seed wing holds the seed tightly in a cup, leaves have primary stomatal bands abaxial (below the phloem vessels) only.
* Subfamily Abietoideae
'', and ''Tsuga
''): cones are annual, without a distinct umbo, the cone scale base is narrow, with the seeds partly visible in abaxial view, the seed has resin vesicles, the seed wing holds the seed tightly in a cup, leaves have primary stomatal bands abaxial (below the phloem vessels) only.
Some genera of interest include ''Pinus
'' and ''Pseudotsuga
.'' The oldest record of the family is ''Eathiestrobus
'' from the Upper Jurassic of Scotland.
A revised 2018 phylogeny places ''Cathaya'' as sister to the pines rather than in the Laricoidae subfamily with ''Larix'' and ''Pseudotsuga''.
External stresses on plants have the ability to change the structure and composition of forest ecosystems
. Common external stress that ''Pinaceae'' experience are herbivore
attack which often leads to tree death.
In order to combat these stresses, trees need to adapt or evolve defenses against these stresses. ''Pinaceae'' have evolved a myriad of mechanical and chemical defenses, or a combination of the two, in order to protect themselves against antagonists.
''Pinaceae'' have the ability to up-regulate a combination of constitutive mechanical and chemical strategies
to further their defenses.
''Pinaceae'' defenses are prevalent in the bark of the trees. This part of the tree contributes a complex defensive boundary against external antagonists.
[Franceschi, V. R., P. Krokene, T. Krekling, and E. Christiansen. 2000. Phloem parenchyma cells are involved in local and distance defense response to fungal inoculation or bark-beetle attack in Norway spruce (''Pinaceae''). American Journal of Botany 87:314-326.] Constitutive
and induced defenses
are both found in the bark.
s are typically the first line of defenses used against antagonists and can include sclerified cells, lignified periderm cells, and secondary compounds such as phenolics
Constitutive defenses are always expressed and offer immediate protection from invaders but could also be defeated by antagonists that have evolved adaptations to these defense mechanisms.
One of the common secondary compounds used by ''Pinaceae'' are phenolics or polyphenols. These secondary compounds are preserved in vacuole
s of polyphenolic parenchyma cells
(PP) in the secondary phloem
responses need to be activated by certain cues, such as herbivore damage or other biotic signals.
A common induced defense mechanism used by ''Pinaceae'' is resins.
s are also one of the primary defenses used against attack.
Resins are short term defenses that are composed of a complex combination of volatile mono
) and sesquiterpene
) and nonvolatile diterpene
resin acids (C20
They are produced and stored in specialized secretory areas known as resin ducts, resin blisters, or resin cavities.
Resins have the ability to wash away, trap, fend off antagonists, and are also involved in wound sealing.
They are an effective defense mechanism because they have toxic and inhibitory effects on invaders, such as insects or pathogens.
Resins could have developed as an evolutionary defense against bark beetle
One well researched resin present in ''Pinaceae'' is oleoresin
. Oleoresin had been found to be a valuable part of the conifer
defense mechanism against biotic attacks
They are found in secretory tissues
in tree stems, roots, and leaves.
Oleoresin is also needed in order to classify conifers.
Active research: methyl jasmonate (MJ)
The topic of defense mechanisms within family ''Pinaceae'' is a very active area of study with numerous studies being conducted. Many of these studies use methyl jasmonate
(MJ) as an antagonist.
Methyl jasmonate is known to be able to induce defense responses in the stems of multiple ''Pinaceae'' species.
It has been found that MJ stimulated the activation of PP cells and formation of xylem traumatic resin ducts (TD). These are structures that are involved in the release of phenolics and resins, both forms of defense mechanism.
File:Pinceae_Bishop_pine_prickle_cone_pine_pinus_muricata.jpg | Close up of bishop pine cones
File:Pinaceae_Knobcone_Pine_Pinus_attenuata.jpg | Knobcone pine cone
Arboretum de Villardebelle
French Arboretum of conifers around the world
Pinaceae on the web page of the Tree-of-Life project40 Pine Trees From Around the World
by The Spruce
Key to Pinaceae
from the Jepson Manual, covers Californian species and much of western North America
Pinaceae in Flora of North AmericaPinus in USDA Plants Database